{"results":[{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"P07339","secondaryAccessions":["Q6IB57"],"uniProtkbId":"CATD_HUMAN","entryAudit":{"firstPublicDate":"1988-04-01","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"1988-04-01","entryVersion":258,"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":"Cathepsin D"},"ecNumbers":[{"value":"3.4.23.5"}]},"contains":[{"recommendedName":{"fullName":{"value":"Cathepsin D light chain"}}},{"recommendedName":{"fullName":{"value":"Cathepsin D heavy chain"}}}],"flag":"Precursor"},"genes":[{"geneName":{"value":"CTSD"},"synonyms":[{"value":"CPSD"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27333034"}],"value":"Acid protease active in intracellular protein breakdown. Plays a role in APP processing following cleavage and activation by ADAM30 which leads to APP degradation (PubMed:27333034). Involved in the pathogenesis of several diseases such as breast cancer and possibly Alzheimer disease"}],"commentType":"FUNCTION"},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5"}},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P18242"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"1426530"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27333034"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8393577"}],"value":"Consists of a light chain and a heavy chain (PubMed:1426530, PubMed:8393577). Interacts with ADAM30; this leads to activation of CTSD (PubMed:27333034). Interacts with GRN; stabilizes CTSD; increases its proteolytic activity (By similarity)"}],"commentType":"SUBUNIT"},{"commentType":"INTERACTION","interactions":[{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P05067","geneName":"APP","intActId":"EBI-77613"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q9P1A6-3","geneName":"DLGAP2","intActId":"EBI-12019838"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"I6L9I8","geneName":"EPN3","intActId":"EBI-12866582"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q9H6S3","geneName":"EPS8L2","intActId":"EBI-3940939"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q7Z602","geneName":"GPR141","intActId":"EBI-21649723"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P28799","geneName":"GRN","intActId":"EBI-747754"},"numberOfExperiments":4,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P28799","geneName":"GRN","chainId":"PRO_0000012695","intActId":"EBI-21335602"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P28799","geneName":"GRN","chainId":"PRO_0000012696","intActId":"EBI-21335615"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P28799","geneName":"GRN","chainId":"PRO_0000012697","intActId":"EBI-21335629"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P28799","geneName":"GRN","chainId":"PRO_0000012698","intActId":"EBI-21335642"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P28799","geneName":"GRN","chainId":"PRO_0000012699","intActId":"EBI-21335656"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P28799","geneName":"GRN","chainId":"PRO_0000012700","intActId":"EBI-21335669"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P28799","geneName":"GRN","chainId":"PRO_0000012701","intActId":"EBI-21335682"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P68431","geneName":"H3C12","intActId":"EBI-79722"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q9Y6F6-3","geneName":"IRAG1","intActId":"EBI-25840037"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q12756","geneName":"KIF1A","intActId":"EBI-2679809"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q5TA79","geneName":"LCE2A","intActId":"EBI-10246607"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q86VF5-3","geneName":"MOGAT3","intActId":"EBI-25840143"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"O15130-2","geneName":"NPFF","intActId":"EBI-25840002"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q96LB9","geneName":"PGLYRP3","intActId":"EBI-12339509"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P09565","geneName":"PP9974","intActId":"EBI-10196507"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q9C004","geneName":"SPRY4","intActId":"EBI-354861"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q8NBJ7","geneName":"SUMF2","intActId":"EBI-723091"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q9BQG1","geneName":"SYT3","intActId":"EBI-17284568"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P28347-2","geneName":"TEAD1","intActId":"EBI-12151837"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"P45880","geneName":"VDAC2","intActId":"EBI-354022"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q15007-2","geneName":"WTAP","intActId":"EBI-25840023"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"O00308","geneName":"WWP2","intActId":"EBI-743923"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q5W0Z9-4","geneName":"ZDHHC20","intActId":"EBI-25840130"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P07339","intActId":"EBI-2115097"},"interactantTwo":{"uniProtKBAccession":"Q6ZNH5","geneName":"ZNF497","intActId":"EBI-10486136"},"numberOfExperiments":4,"organismDiffer":false}]},{"commentType":"SUBCELLULAR LOCATION","note":{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20551380"}],"value":"Identified by mass spectrometry in melanosome fractions from stage I to stage IV. In aortic samples, detected as an extracellular protein loosely bound to the matrix (PubMed:20551380)"}]},"subcellularLocations":[{"location":{"value":"Lysosome","id":"SL-0158"}},{"location":{"value":"Melanosome","id":"SL-0161"}},{"location":{"value":"Secreted, extracellular space","id":"SL-0112"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"1426530"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20551380"}],"value":"Expressed in the aorta extracellular space (at protein level) (PubMed:20551380). Expressed in liver (at protein level) (PubMed:1426530)"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12754519"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16263699"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16335952"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19159218"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23234360"}],"value":"N- and O-glycosylated"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27333034"}],"value":"Undergoes proteolytic cleavage and activation by ADAM30"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"1426530"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27333034"}],"value":"As well as the major heavy chain which starts at Leu-169, 2 minor forms starting at Gly-170 and Gly-171 have been identified (PubMed:1426530). An additional form starting at Ala-168 has also been identified (PubMed:27333034)"}],"commentType":"PTM"},{"texts":[{"value":"The Val-58 allele is significantly overrepresented in demented patients (11.8%) compared with non-demented controls (4.9%). Carriers of the Val-58 allele have a 3.1-fold increased risk for developing AD than non-carriers"}],"commentType":"POLYMORPHISM"},{"commentType":"DISEASE","disease":{"diseaseId":"Ceroid lipofuscinosis, neuronal, 10","diseaseAccession":"DI-00818","acronym":"CLN10","description":"A form of neuronal ceroid lipofuscinosis with onset at birth or early childhood. Neuronal ceroid lipofuscinoses are progressive neurodegenerative, lysosomal storage diseases characterized by intracellular accumulation of autofluorescent liposomal material, and clinically by seizures, dementia, visual loss, and/or cerebral atrophy.","diseaseCrossReference":{"database":"MIM","id":"610127"},"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16670177"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16685649"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21990111"}]},"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 peptidase A1 family"}],"commentType":"SIMILARITY"},{"commentType":"WEB RESOURCE","resourceName":"NCL CTSD","resourceUrl":"https://www.ucl.ac.uk/ncl-disease/ncl-resource-gateway-batten-disease","ftp":false,"note":"Neural Ceroid Lipofuscinoses mutation db"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000255"}]},{"type":"Propeptide","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":64,"modifier":"EXACT"}},"description":"Activation peptide","featureId":"PRO_0000025949"},{"type":"Chain","location":{"start":{"value":65,"modifier":"EXACT"},"end":{"value":412,"modifier":"EXACT"}},"description":"Cathepsin D","featureId":"PRO_0000025950"},{"type":"Chain","location":{"start":{"value":65,"modifier":"EXACT"},"end":{"value":162,"modifier":"EXACT"}},"description":"Cathepsin D light chain","evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"1426530"}],"featureId":"PRO_0000025951"},{"type":"Chain","location":{"start":{"value":169,"modifier":"EXACT"},"end":{"value":412,"modifier":"EXACT"}},"description":"Cathepsin D heavy chain","evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"1426530"}],"featureId":"PRO_0000025952"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":407,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000255","source":"PROSITE-ProRule","id":"PRU01103"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000255","source":"PROSITE-ProRule","id":"PRU10094"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8393577"}]},{"type":"Active site","location":{"start":{"value":295,"modifier":"EXACT"},"end":{"value":295,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000255","source":"PROSITE-ProRule","id":"PRU10094"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8393577"}]},{"type":"Glycosylation","location":{"start":{"value":63,"modifier":"EXACT"},"end":{"value":63,"modifier":"EXACT"}},"description":"O-linked (GalNAc...) threonine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23234360"}],"featureId":""},{"type":"Glycosylation","location":{"start":{"value":134,"modifier":"EXACT"},"end":{"value":134,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19159218"}],"featureId":""},{"type":"Glycosylation","location":{"start":{"value":263,"modifier":"EXACT"},"end":{"value":263,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12754519"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16263699"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16335952"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19159218"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8393577"}],"featureId":""},{"type":"Disulfide bond","location":{"start":{"value":91,"modifier":"EXACT"},"end":{"value":160,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8393577"}]},{"type":"Disulfide bond","location":{"start":{"value":110,"modifier":"EXACT"},"end":{"value":117,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8393577"}]},{"type":"Disulfide bond","location":{"start":{"value":286,"modifier":"EXACT"},"end":{"value":290,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8393577"}]},{"type":"Disulfide bond","location":{"start":{"value":329,"modifier":"EXACT"},"end":{"value":366,"modifier":"EXACT"}},"description":""},{"type":"Natural variant","location":{"start":{"value":58,"modifier":"EXACT"},"end":{"value":58,"modifier":"EXACT"}},"description":"associated with increased risk for AD; possibly influences secretion and intracellular maturation; dbSNP:rs17571","featureCrossReferences":[{"database":"dbSNP","id":"rs17571"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10716266"}],"featureId":"VAR_011621","alternativeSequence":{"originalSequence":"A","alternativeSequences":["V"]}},{"type":"Natural variant","location":{"start":{"value":229,"modifier":"EXACT"},"end":{"value":229,"modifier":"EXACT"}},"description":"in CLN10; dbSNP:rs121912789","featureCrossReferences":[{"database":"dbSNP","id":"rs121912789"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16685649"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21990111"}],"featureId":"VAR_029362","alternativeSequence":{"originalSequence":"F","alternativeSequences":["I"]}},{"type":"Natural variant","location":{"start":{"value":282,"modifier":"EXACT"},"end":{"value":282,"modifier":"EXACT"}},"description":"in dbSNP:rs147278302","featureCrossReferences":[{"database":"dbSNP","id":"rs147278302"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16670177"}],"featureId":"VAR_058490","alternativeSequence":{"originalSequence":"G","alternativeSequences":["R"]}},{"type":"Natural variant","location":{"start":{"value":383,"modifier":"EXACT"},"end":{"value":383,"modifier":"EXACT"}},"description":"in CLN10; dbSNP:rs121912790","featureCrossReferences":[{"database":"dbSNP","id":"rs121912790"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16685649"}],"featureId":"VAR_029363","alternativeSequence":{"originalSequence":"W","alternativeSequences":["C"]}},{"type":"Beta strand","location":{"start":{"value":67,"modifier":"EXACT"},"end":{"value":74,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Turn","location":{"start":{"value":75,"modifier":"EXACT"},"end":{"value":77,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":78,"modifier":"EXACT"},"end":{"value":85,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Turn","location":{"start":{"value":86,"modifier":"EXACT"},"end":{"value":89,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":90,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":103,"modifier":"EXACT"},"end":{"value":107,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":115,"modifier":"EXACT"},"end":{"value":118,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":125,"modifier":"EXACT"},"end":{"value":127,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":132,"modifier":"EXACT"},"end":{"value":141,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":146,"modifier":"EXACT"},"end":{"value":159,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":172,"modifier":"EXACT"},"end":{"value":184,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Turn","location":{"start":{"value":189,"modifier":"EXACT"},"end":{"value":191,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":196,"modifier":"EXACT"},"end":{"value":200,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":204,"modifier":"EXACT"},"end":{"value":206,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":208,"modifier":"EXACT"},"end":{"value":210,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":211,"modifier":"EXACT"},"end":{"value":213,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":214,"modifier":"EXACT"},"end":{"value":220,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":224,"modifier":"EXACT"},"end":{"value":226,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QBH"}]},{"type":"Beta strand","location":{"start":{"value":228,"modifier":"EXACT"},"end":{"value":233,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":236,"modifier":"EXACT"},"end":{"value":238,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"1LYW"}]},{"type":"Beta strand","location":{"start":{"value":243,"modifier":"EXACT"},"end":{"value":247,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":252,"modifier":"EXACT"},"end":{"value":254,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":255,"modifier":"EXACT"},"end":{"value":263,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":265,"modifier":"EXACT"},"end":{"value":267,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":270,"modifier":"EXACT"},"end":{"value":274,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":276,"modifier":"EXACT"},"end":{"value":279,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Turn","location":{"start":{"value":280,"modifier":"EXACT"},"end":{"value":282,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"1LYW"}]},{"type":"Beta strand","location":{"start":{"value":284,"modifier":"EXACT"},"end":{"value":286,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":290,"modifier":"EXACT"},"end":{"value":294,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":299,"modifier":"EXACT"},"end":{"value":303,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":305,"modifier":"EXACT"},"end":{"value":315,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":318,"modifier":"EXACT"},"end":{"value":321,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"1LYA"}]},{"type":"Beta strand","location":{"start":{"value":325,"modifier":"EXACT"},"end":{"value":327,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":329,"modifier":"EXACT"},"end":{"value":334,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":335,"modifier":"EXACT"},"end":{"value":337,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"4OBZ"}]},{"type":"Beta strand","location":{"start":{"value":338,"modifier":"EXACT"},"end":{"value":342,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":345,"modifier":"EXACT"},"end":{"value":349,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":351,"modifier":"EXACT"},"end":{"value":354,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":355,"modifier":"EXACT"},"end":{"value":360,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":363,"modifier":"EXACT"},"end":{"value":372,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":374,"modifier":"EXACT"},"end":{"value":376,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Turn","location":{"start":{"value":377,"modifier":"EXACT"},"end":{"value":379,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":383,"modifier":"EXACT"},"end":{"value":385,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Helix","location":{"start":{"value":387,"modifier":"EXACT"},"end":{"value":392,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":393,"modifier":"EXACT"},"end":{"value":398,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Turn","location":{"start":{"value":399,"modifier":"EXACT"},"end":{"value":402,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]},{"type":"Beta strand","location":{"start":{"value":403,"modifier":"EXACT"},"end":{"value":409,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0007829","source":"PDB","id":"6QCB"}]}],"keywords":[{"id":"KW-0002","category":"Technical term","name":"3D-structure"},{"id":"KW-0026","category":"Disease","name":"Alzheimer disease"},{"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"id":"KW-0903","category":"Technical term","name":"Direct protein sequencing"},{"id":"KW-0225","category":"Disease","name":"Disease variant"},{"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"id":"KW-0523","category":"Disease","name":"Neurodegeneration"},{"id":"KW-0525","category":"Disease","name":"Neuronal ceroid lipofuscinosis"},{"id":"KW-0645","category":"Molecular function","name":"Protease"},{"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"id":"KW-0964","category":"Cellular component","name":"Secreted"},{"id":"KW-0732","category":"Domain","name":"Signal"},{"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"3927292","citationType":"journal article","authors":["Faust P.L.","Kornfeld S.","Chirgwin J.M."],"citationCrossReferences":[{"database":"PubMed","id":"3927292"},{"database":"DOI","id":"10.1073/pnas.82.15.4910"}],"title":"Cloning and sequence analysis of cDNA for human cathepsin D.","publicationDate":"1985","journal":"Proc. Natl. Acad. Sci. U.S.A.","firstPage":"4910","lastPage":"4914","volume":"82"},"referencePositions":["NUCLEOTIDE SEQUENCE [MRNA]"]},{"referenceNumber":2,"citation":{"id":"3588310","citationType":"journal article","authors":["Westley B.R.","May F.E.B."],"citationCrossReferences":[{"database":"PubMed","id":"3588310"},{"database":"DOI","id":"10.1093/nar/15.9.3773"}],"title":"Oestrogen regulates cathepsin D mRNA levels in oestrogen responsive human breast cancer cells.","publicationDate":"1987","journal":"Nucleic Acids Res.","firstPage":"3773","lastPage":"3786","volume":"15"},"referencePositions":["NUCLEOTIDE SEQUENCE [MRNA]"]},{"referenceNumber":3,"citation":{"id":"2069717","citationType":"journal article","authors":["Redecker B.","Heckendorf B.","Grosch H.W.","Mersmann G.","Hasilik A."],"citationCrossReferences":[{"database":"PubMed","id":"2069717"},{"database":"DOI","id":"10.1089/dna.1991.10.423"}],"title":"Molecular organization of the human cathepsin D gene.","publicationDate":"1991","journal":"DNA Cell Biol.","firstPage":"423","lastPage":"431","volume":"10"},"referencePositions":["NUCLEOTIDE SEQUENCE [GENOMIC DNA]"]},{"referenceNumber":4,"citation":{"id":"CI-D49P6QQO8F5LU","citationType":"submission","authors":["Ebert L.","Schick M.","Neubert P.","Schatten R.","Henze S.","Korn B."],"title":"Cloning of human full open reading frames in Gateway(TM) system entry vector (pDONR201).","publicationDate":"JUN-2004","submissionDatabase":"EMBL/GenBank/DDBJ databases"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA]"]},{"referenceNumber":5,"citation":{"id":"CI-54UTT6AFSFO1R","citationType":"submission","authors":["Kalnine N.","Chen X.","Rolfs A.","Halleck A.","Hines L.","Eisenstein S.","Koundinya M.","Raphael J.","Moreira D.","Kelley T.","LaBaer J.","Lin Y.","Phelan M.","Farmer A."],"title":"Cloning of human full-length CDSs in BD Creator(TM) system donor vector.","publicationDate":"OCT-2004","submissionDatabase":"EMBL/GenBank/DDBJ databases"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA]"]},{"referenceNumber":6,"citation":{"id":"15489334","citationType":"journal article","authoringGroup":["The MGC Project Team"],"citationCrossReferences":[{"database":"PubMed","id":"15489334"},{"database":"DOI","id":"10.1101/gr.2596504"}],"title":"The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC).","publicationDate":"2004","journal":"Genome Res.","firstPage":"2121","lastPage":"2127","volume":"14"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA]"],"referenceComments":[{"value":"Kidney","type":"TISSUE"}]},{"referenceNumber":7,"citation":{"id":"8262386","citationType":"journal article","authors":["May F.E.","Smith D.J.","Westley B.R."],"citationCrossReferences":[{"database":"PubMed","id":"8262386"},{"database":"DOI","id":"10.1016/0378-1119(93)90107-e"}],"title":"The human cathepsin D-encoding gene is transcribed from an estrogen-regulated and a constitutive start point.","publicationDate":"1993","journal":"Gene","firstPage":"277","lastPage":"282","volume":"134"},"referencePositions":["NUCLEOTIDE SEQUENCE [GENOMIC DNA] OF 1-22"]},{"referenceNumber":8,"citation":{"id":"7935485","citationType":"journal article","authors":["Augereau P.","Miralles F.","Cavailles V.","Gaudelet C.","Parker M.","Rochefort H."],"citationCrossReferences":[{"database":"PubMed","id":"7935485"},{"database":"DOI","id":"10.1210/mend.8.6.7935485"}],"title":"Characterization of the proximal estrogen-responsive element of human cathepsin D gene.","publicationDate":"1994","journal":"Mol. Endocrinol.","firstPage":"693","lastPage":"703","volume":"8"},"referencePositions":["NUCLEOTIDE SEQUENCE [GENOMIC DNA] OF 1-22"]},{"referenceNumber":9,"citation":{"id":"CI-6U26R306QNDO8","citationType":"submission","authors":["Hochstrasser D.F.","Frutiger S.","Paquet N.","Bairoch A.","Ravier F.","Pasquali C.","Sanchez J.-C.","Tissot J.-D.","Bjellqvist B.","Vargas R.","Appel R.D.","Hughes G.J."],"publicationDate":"JUN-1992","submissionDatabase":"UniProtKB"},"referencePositions":["PROTEIN SEQUENCE OF 170-180"],"referenceComments":[{"value":"Liver","type":"TISSUE"}]},{"referenceNumber":10,"citation":{"id":"1426530","citationType":"journal article","authors":["Kobayashi T.","Honke K.","Gasa S.","Fujii T.","Maguchi S.","Miyazaki T.","Makita A."],"citationCrossReferences":[{"database":"PubMed","id":"1426530"},{"database":"DOI","id":"10.1016/0020-711x(92)90076-d"}],"title":"Proteolytic processing sites producing the mature form of human cathepsin D.","publicationDate":"1992","journal":"Int. J. Biochem.","firstPage":"1487","lastPage":"1491","volume":"24"},"referencePositions":["PROTEIN SEQUENCE OF 154-162 AND 169-180","SUBUNIT","TISSUE SPECIFICITY"]},{"referenceNumber":11,"citation":{"id":"12643545","citationType":"journal article","authors":["Basrur V.","Yang F.","Kushimoto T.","Higashimoto Y.","Yasumoto K.","Valencia J.","Muller J.","Vieira W.D.","Watabe H.","Shabanowitz J.","Hearing V.J.","Hunt D.F.","Appella E."],"citationCrossReferences":[{"database":"PubMed","id":"12643545"},{"database":"DOI","id":"10.1021/pr025562r"}],"title":"Proteomic analysis of early melanosomes: identification of novel melanosomal proteins.","publicationDate":"2003","journal":"J. Proteome Res.","firstPage":"69","lastPage":"79","volume":"2"},"referencePositions":["SUBCELLULAR LOCATION [LARGE SCALE ANALYSIS]"],"referenceComments":[{"value":"Melanoma","type":"TISSUE"}]},{"referenceNumber":12,"citation":{"id":"12754519","citationType":"journal article","authors":["Zhang H.","Li X.-J.","Martin D.B.","Aebersold R."],"citationCrossReferences":[{"database":"PubMed","id":"12754519"},{"database":"DOI","id":"10.1038/nbt827"}],"title":"Identification and quantification of N-linked glycoproteins using hydrazide chemistry, stable isotope labeling and mass spectrometry.","publicationDate":"2003","journal":"Nat. Biotechnol.","firstPage":"660","lastPage":"666","volume":"21"},"referencePositions":["GLYCOSYLATION AT ASN-263"]},{"referenceNumber":13,"citation":{"id":"16335952","citationType":"journal article","authors":["Liu T.","Qian W.-J.","Gritsenko M.A.","Camp D.G. II","Monroe M.E.","Moore R.J.","Smith R.D."],"citationCrossReferences":[{"database":"PubMed","id":"16335952"},{"database":"DOI","id":"10.1021/pr0502065"}],"title":"Human plasma N-glycoproteome analysis by immunoaffinity subtraction, hydrazide chemistry, and mass spectrometry.","publicationDate":"2005","journal":"J. Proteome Res.","firstPage":"2070","lastPage":"2080","volume":"4"},"referencePositions":["GLYCOSYLATION [LARGE SCALE ANALYSIS] AT ASN-263"],"referenceComments":[{"value":"Plasma","type":"TISSUE"}]},{"referenceNumber":14,"citation":{"id":"16670177","citationType":"journal article","authors":["Siintola E.","Partanen S.","Stromme P.","Haapanen A.","Haltia M.","Maehlen J.","Lehesjoki A.E.","Tyynela J."],"citationCrossReferences":[{"database":"PubMed","id":"16670177"},{"database":"DOI","id":"10.1093/brain/awl107"}],"title":"Cathepsin D deficiency underlies congenital human neuronal ceroid-lipofuscinosis.","publicationDate":"2006","journal":"Brain","firstPage":"1438","lastPage":"1445","volume":"129"},"referencePositions":["INVOLVEMENT IN CLN10","VARIANT ARG-282"]},{"referenceNumber":15,"citation":{"id":"17081065","citationType":"journal article","authors":["Chi A.","Valencia J.C.","Hu Z.-Z.","Watabe H.","Yamaguchi H.","Mangini N.J.","Huang H.","Canfield V.A.","Cheng K.C.","Yang F.","Abe R.","Yamagishi S.","Shabanowitz J.","Hearing V.J.","Wu C.","Appella E.","Hunt D.F."],"citationCrossReferences":[{"database":"PubMed","id":"17081065"},{"database":"DOI","id":"10.1021/pr060363j"}],"title":"Proteomic and bioinformatic characterization of the biogenesis and function of melanosomes.","publicationDate":"2006","journal":"J. Proteome Res.","firstPage":"3135","lastPage":"3144","volume":"5"},"referencePositions":["SUBCELLULAR LOCATION [LARGE SCALE ANALYSIS]"],"referenceComments":[{"value":"Melanoma","type":"TISSUE"}]},{"referenceNumber":16,"citation":{"id":"16263699","citationType":"journal article","authors":["Lewandrowski U.","Moebius J.","Walter U.","Sickmann A."],"citationCrossReferences":[{"database":"PubMed","id":"16263699"},{"database":"DOI","id":"10.1074/mcp.m500324-mcp200"}],"title":"Elucidation of N-glycosylation sites on human platelet proteins: a glycoproteomic approach.","publicationDate":"2006","journal":"Mol. Cell. Proteomics","firstPage":"226","lastPage":"233","volume":"5"},"referencePositions":["GLYCOSYLATION [LARGE SCALE ANALYSIS] AT ASN-263"],"referenceComments":[{"value":"Platelet","type":"TISSUE"}]},{"referenceNumber":17,"citation":{"id":"19159218","citationType":"journal article","authors":["Chen R.","Jiang X.","Sun D.","Han G.","Wang F.","Ye M.","Wang L.","Zou H."],"citationCrossReferences":[{"database":"PubMed","id":"19159218"},{"database":"DOI","id":"10.1021/pr8008012"}],"title":"Glycoproteomics analysis of human liver tissue by combination of multiple enzyme digestion and hydrazide chemistry.","publicationDate":"2009","journal":"J. Proteome Res.","firstPage":"651","lastPage":"661","volume":"8"},"referencePositions":["GLYCOSYLATION [LARGE SCALE ANALYSIS] AT ASN-134 AND ASN-263"],"referenceComments":[{"value":"Liver","type":"TISSUE"}]},{"referenceNumber":18,"citation":{"id":"20551380","citationType":"journal article","authors":["Didangelos A.","Yin X.","Mandal K.","Baumert M.","Jahangiri M.","Mayr M."],"citationCrossReferences":[{"database":"PubMed","id":"20551380"},{"database":"DOI","id":"10.1074/mcp.m110.001693"}],"title":"Proteomics characterization of extracellular space components in the human aorta.","publicationDate":"2010","journal":"Mol. Cell. Proteomics","firstPage":"2048","lastPage":"2062","volume":"9"},"referencePositions":["TISSUE SPECIFICITY","SUBCELLULAR LOCATION"]},{"referenceNumber":19,"citation":{"id":"21269460","citationType":"journal article","authors":["Burkard T.R.","Planyavsky M.","Kaupe I.","Breitwieser F.P.","Buerckstuemmer T.","Bennett K.L.","Superti-Furga G.","Colinge J."],"citationCrossReferences":[{"database":"PubMed","id":"21269460"},{"database":"DOI","id":"10.1186/1752-0509-5-17"}],"title":"Initial characterization of the human central proteome.","publicationDate":"2011","journal":"BMC Syst. Biol.","firstPage":"17","lastPage":"17","volume":"5"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"]},{"referenceNumber":20,"citation":{"id":"23234360","citationType":"journal article","authors":["Halim A.","Ruetschi U.","Larson G.","Nilsson J."],"citationCrossReferences":[{"database":"PubMed","id":"23234360"},{"database":"DOI","id":"10.1021/pr300963h"}],"title":"LC-MS/MS characterization of O-glycosylation sites and glycan structures of human cerebrospinal fluid glycoproteins.","publicationDate":"2013","journal":"J. Proteome Res.","firstPage":"573","lastPage":"584","volume":"12"},"referencePositions":["GLYCOSYLATION AT THR-63","IDENTIFICATION BY MASS SPECTROMETRY"]},{"referenceNumber":21,"citation":{"id":"25944712","citationType":"journal article","authors":["Vaca Jacome A.S.","Rabilloud T.","Schaeffer-Reiss C.","Rompais M.","Ayoub D.","Lane L.","Bairoch A.","Van Dorsselaer A.","Carapito C."],"citationCrossReferences":[{"database":"PubMed","id":"25944712"},{"database":"DOI","id":"10.1002/pmic.201400617"}],"title":"N-terminome analysis of the human mitochondrial proteome.","publicationDate":"2015","journal":"Proteomics","firstPage":"2519","lastPage":"2524","volume":"15"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"]},{"referenceNumber":22,"citation":{"id":"27333034","citationType":"journal article","authors":["Letronne F.","Laumet G.","Ayral A.M.","Chapuis J.","Demiautte F.","Laga M.","Vandenberghe M.E.","Malmanche N.","Leroux F.","Eysert F.","Sottejeau Y.","Chami L.","Flaig A.","Bauer C.","Dourlen P.","Lesaffre M.","Delay C.","Huot L.","Dumont J.","Werkmeister E.","Lafont F.","Mendes T.","Hansmannel F.","Dermaut B.","Deprez B.","Herard A.S.","Dhenain M.","Souedet N.","Pasquier F.","Tulasne D.","Berr C.","Hauw J.J.","Lemoine Y.","Amouyel P.","Mann D.","Deprez R.","Checler F.","Hot D.","Delzescaux T.","Gevaert K.","Lambert J.C."],"citationCrossReferences":[{"database":"PubMed","id":"27333034"},{"database":"DOI","id":"10.1016/j.ebiom.2016.06.002"}],"title":"ADAM30 Downregulates APP-Linked Defects Through Cathepsin D Activation in Alzheimer's Disease.","publicationDate":"2016","journal":"EBioMedicine","firstPage":"278","lastPage":"292","volume":"9"},"referencePositions":["INTERACTION WITH ADAM30","IDENTIFICATION BY MASS SPECTROMETRY","CLEAVAGE"]},{"referenceNumber":23,"citation":{"id":"8467789","citationType":"journal article","authors":["Metcalf P.","Fusek M."],"citationCrossReferences":[{"database":"PubMed","id":"8467789"},{"database":"DOI","id":"10.1002/j.1460-2075.1993.tb05774.x"}],"title":"Two crystal structures for cathepsin D: the lysosomal targeting signal and active site.","publicationDate":"1993","journal":"EMBO J.","firstPage":"1293","lastPage":"1302","volume":"12"},"referencePositions":["X-RAY CRYSTALLOGRAPHY (3 ANGSTROMS)"],"referenceComments":[{"value":"Spleen","type":"TISSUE"}]},{"referenceNumber":24,"citation":{"id":"8393577","citationType":"journal article","authors":["Baldwin E.T.","Bhat T.N.","Gulnik S.","Hosur M.V.","Sowder R.C. II","Cachau R.E.","Collins J.","Silva A.M.","Erickson J.W."],"citationCrossReferences":[{"database":"PubMed","id":"8393577"},{"database":"DOI","id":"10.1073/pnas.90.14.6796"}],"title":"Crystal structures of native and inhibited forms of human cathepsin D: implications for lysosomal targeting and drug design.","publicationDate":"1993","journal":"Proc. Natl. Acad. Sci. U.S.A.","firstPage":"6796","lastPage":"6800","volume":"90"},"referencePositions":["X-RAY CRYSTALLOGRAPHY (2.5 ANGSTROMS)","SUBUNIT"],"referenceComments":[{"value":"Liver","type":"TISSUE"}]},{"referenceNumber":25,"citation":{"id":"10716266","citationType":"journal article","authors":["Papassotiropoulos A.","Bagli M.","Kurz A.","Kornhuber J.","Forstl H.","Maier W.","Pauls J.","Lautenschlager N.","Heun R."],"citationCrossReferences":[{"database":"PubMed","id":"10716266"},{"database":"DOI","id":"10.1002/1531-8249(200003)47:3<399::aid-ana22>3.3.co;2-x"}],"title":"A genetic variation of cathepsin D is a major risk factor for Alzheimer's disease.","publicationDate":"2000","journal":"Ann. Neurol.","firstPage":"399","lastPage":"403","volume":"47"},"referencePositions":["VARIANT VAL-58"]},{"referenceNumber":26,"citation":{"id":"16685649","citationType":"journal article","authors":["Steinfeld R.","Reinhardt K.","Schreiber K.","Hillebrand M.","Kraetzner R.","Bruck W.","Saftig P.","Gartner J."],"citationCrossReferences":[{"database":"PubMed","id":"16685649"},{"database":"DOI","id":"10.1086/504159"}],"title":"Cathepsin D deficiency is associated with a human neurodegenerative disorder.","publicationDate":"2006","journal":"Am. J. Hum. Genet.","firstPage":"988","lastPage":"998","volume":"78"},"referencePositions":["VARIANTS CLN10 ILE-229 AND CYS-383"]},{"referenceNumber":27,"citation":{"id":"21990111","citationType":"journal article","authors":["Kousi M.","Lehesjoki A.E.","Mole S.E."],"citationCrossReferences":[{"database":"PubMed","id":"21990111"},{"database":"DOI","id":"10.1002/humu.21624"}],"title":"Update of the mutation spectrum and clinical correlations of over 360 mutations in eight genes that underlie the neuronal ceroid lipofuscinoses.","publicationDate":"2012","journal":"Hum. Mutat.","firstPage":"42","lastPage":"63","volume":"33"},"referencePositions":["VARIANT CLN10 ILE-229"]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"M11233","properties":[{"key":"ProteinId","value":"AAB59529.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"mRNA"}]},{"database":"EMBL","id":"X05344","properties":[{"key":"ProteinId","value":"CAA28955.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"mRNA"}]},{"database":"EMBL","id":"M63138","properties":[{"key":"ProteinId","value":"AAA51922.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"EMBL","id":"M63134","properties":[{"key":"ProteinId","value":"AAA51922.1"},{"key":"Status","value":"JOINED"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"EMBL","id":"M63135","properties":[{"key":"ProteinId","value":"AAA51922.1"},{"key":"Status","value":"JOINED"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"EMBL","id":"M63136","properties":[{"key":"ProteinId","value":"AAA51922.1"},{"key":"Status","value":"JOINED"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"EMBL","id":"M63137","properties":[{"key":"ProteinId","value":"AAA51922.1"},{"key":"Status","value":"JOINED"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"EMBL","id":"CR456947","properties":[{"key":"ProteinId","value":"CAG33228.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"mRNA"}]},{"database":"EMBL","id":"BT006910","properties":[{"key":"ProteinId","value":"AAP35556.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"mRNA"}]},{"database":"EMBL","id":"BT020155","properties":[{"key":"ProteinId","value":"AAV38957.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"mRNA"}]},{"database":"EMBL","id":"BC016320","properties":[{"key":"ProteinId","value":"AAH16320.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"mRNA"}]},{"database":"EMBL","id":"L12980","properties":[{"key":"ProteinId","value":"AAA16314.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"EMBL","id":"S74689","properties":[{"key":"ProteinId","value":"AAD14156.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"EMBL","id":"S52557","properties":[{"key":"ProteinId","value":"AAD13868.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"CCDS","id":"CCDS7725.1","properties":[{"key":"Description","value":"-"}]},{"database":"PIR","id":"A25771","properties":[{"key":"EntryName","value":"KHHUD"}]},{"database":"RefSeq","id":"NP_001900.1","properties":[{"key":"NucleotideSequenceId","value":"NM_001909.5"}]},{"database":"PDB","id":"1LYA","properties":[{"key":"Method","value":"X-ray"},{"key":"Resolution","value":"2.50 A"},{"key":"Chains","value":"A/C=65-161, B/D=170-410"}]},{"database":"PDB","id":"1LYB","properties":[{"key":"Method","value":"X-ray"},{"key":"Resolution","value":"2.50 A"},{"key":"Chains","value":"A/C=65-161, B/D=170-410"}]},{"database":"PDB","id":"1LYW","properties":[{"key":"Method","value":"X-ray"},{"key":"Resolution","value":"2.50 A"},{"key":"Chains","value":"A/C/E/G=65-161, B/D/F/H=170-410"}]},{"database":"PDB","id":"4OBZ","properties":[{"key":"Method","value":"X-ray"},{"key":"Resolution","value":"2.90 A"},{"key":"Chains","value":"A/C=60-162, B/D=170-412"}]},{"database":"PDB","id":"4OC6","properties":[{"key":"Method","value":"X-ray"},{"key":"Resolution","value":"2.64 A"},{"key":"Chains","value":"A=60-162, B=170-412"}]},{"database":"PDB","id":"4OD9","properties":[{"key":"Method","value":"X-ray"},{"key":"Resolution","value":"1.90 A"},{"key":"Chains","value":"A/C=60-162, B/D=170-412"}]},{"database":"PDB","id":"6QBG","properties":[{"key":"Method","value":"X-ray"},{"key":"Resolution","value":"1.80 A"},{"key":"Chains","value":"A=65-162, B=171-411"}]},{"database":"PDB","id":"6QBH","properties":[{"key":"Method","value":"X-ray"},{"key":"Resolution","value":"1.85 A"},{"key":"Chains","value":"A=65-161, B=171-411"}]},{"database":"PDB","id":"6QCB","properties":[{"key":"Method","value":"X-ray"},{"key":"Resolution","value":"1.55 A"},{"key":"Chains","value":"A=65-161, B=171-411"}]},{"database":"PDBsum","id":"1LYA","properties":[{"key":"Description","value":"-"}]},{"database":"PDBsum","id":"1LYB","properties":[{"key":"Description","value":"-"}]},{"database":"PDBsum","id":"1LYW","properties":[{"key":"Description","value":"-"}]},{"database":"PDBsum","id":"4OBZ","properties":[{"key":"Description","value":"-"}]},{"database":"PDBsum","id":"4OC6","properties":[{"key":"Description","value":"-"}]},{"database":"PDBsum","id":"4OD9","properties":[{"key":"Description","value":"-"}]},{"database":"PDBsum","id":"6QBG","properties":[{"key":"Description","value":"-"}]},{"database":"PDBsum","id":"6QBH","properties":[{"key":"Description","value":"-"}]},{"database":"PDBsum","id":"6QCB","properties":[{"key":"Description","value":"-"}]},{"database":"AlphaFoldDB","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"MDposit","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"MDRepo","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"BioGRID","id":"107889","properties":[{"key":"Interactions","value":"177"}]},{"database":"DIP","id":"DIP-43906N","properties":[{"key":"Description","value":"-"}]},{"database":"FunCoup","id":"P07339","properties":[{"key":"Number of interactors","value":"1372"}]},{"database":"IntAct","id":"P07339","properties":[{"key":"Interactions","value":"88"}]},{"database":"MINT","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"NDEx","id":"IQUERY-CP-CTSD","properties":[{"key":"Evidences","value":"7 NDEx IQuery Curated Pathways"}]},{"database":"STRING","id":"9606.ENSP00000236671","properties":[{"key":"Description","value":"-"}]},{"database":"BindingDB","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"ChEMBL","id":"CHEMBL2581","properties":[{"key":"Description","value":"-"}]},{"database":"DrugBank","id":"DB03028","properties":[{"key":"GenericName","value":"1h-Benoximidazole-2-Carboxylic Acid"}]},{"database":"DrugBank","id":"DB03096","properties":[{"key":"GenericName","value":"2-Morpholinoethylamine"}]},{"database":"DrugBank","id":"DB07542","properties":[{"key":"GenericName","value":"5-Amino-6-cyclohexyl-4-hydroxy-2-isobutyl-hexanoic acid"}]},{"database":"DrugBank","id":"DB08740","properties":[{"key":"GenericName","value":"CYCLOHEXYLMETHYL-2,3-DIHYDROXY-5-METHYL-HEXYLAMIDE"}]},{"database":"DrugBank","id":"DB02216","properties":[{"key":"GenericName","value":"S-Methylcysteine"}]},{"database":"DrugCentral","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"GuidetoPHARMACOLOGY","id":"2345","properties":[{"key":"Description","value":"-"}]},{"database":"MEROPS","id":"A01.009","properties":[{"key":"Description","value":"-"}]},{"database":"GlyConnect","id":"P07339","properties":[{"key":"glycosylation","value":"45 N-Linked glycans (2 sites)"}]},{"database":"GlyCosmos","id":"P07339","properties":[{"key":"glycosylation","value":"3 sites, 44 glycans"}]},{"database":"GlyGen","id":"P07339","properties":[{"key":"glycosylation","value":"12 sites, 113 N-linked glycans (2 sites), 1 N-linked;o-linked glycan (1 site), 3 O-linked glycans (8 sites)"}]},{"database":"iPTMnet","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"MetOSite","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"PhosphoSitePlus","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"SwissPalm","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"BioMuta","id":"CTSD","properties":[{"key":"Description","value":"-"}]},{"database":"DMDM","id":"115717","properties":[{"key":"Description","value":"-"}]},{"database":"REPRODUCTION-2DPAGE","id":"IPI00011229","properties":[{"key":"Description","value":"-"}]},{"database":"CPTAC","id":"CPTAC-486","properties":[{"key":"Description","value":"-"}]},{"database":"CPTAC","id":"CPTAC-487","properties":[{"key":"Description","value":"-"}]},{"database":"CPTAC","id":"CPTAC-658","properties":[{"key":"Description","value":"-"}]},{"database":"jPOST","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"MassIVE","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"PaxDb","id":"9606-ENSP00000236671","properties":[{"key":"Description","value":"-"}]},{"database":"PeptideAtlas","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"ProteomicsDB","id":"51994","properties":[{"key":"Description","value":"-"}]},{"database":"Pumba","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"TopDownProteomics","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"Antibodypedia","id":"880","properties":[{"key":"antibodies","value":"1212 antibodies from 49 providers"}]},{"database":"DNASU","id":"1509","properties":[{"key":"Description","value":"-"}]},{"database":"Ensembl","id":"ENST00000236671.7","properties":[{"key":"ProteinId","value":"ENSP00000236671.2"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"GeneID","id":"1509","properties":[{"key":"Description","value":"-"}]},{"database":"KEGG","id":"hsa:1509","properties":[{"key":"Description","value":"-"}]},{"database":"MANE-Select","id":"ENST00000236671.7","properties":[{"key":"ProteinId","value":"ENSP00000236671.2"},{"key":"RefSeqNucleotideId","value":"NM_001909.5"},{"key":"RefSeqProteinId","value":"NP_001900.1"}]},{"database":"AGR","id":"HGNC:2529","properties":[{"key":"Description","value":"-"}]},{"database":"ClinPGx","id":"PA27029","properties":[{"key":"Description","value":"-"}]},{"database":"CTD","id":"1509","properties":[{"key":"Description","value":"-"}]},{"database":"DisGeNET","id":"1509","properties":[{"key":"Description","value":"-"}]},{"database":"GeneCards","id":"CTSD","properties":[{"key":"Description","value":"-"}]},{"database":"GeneReviews","id":"CTSD","properties":[{"key":"Description","value":"-"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"HPA","id":"ENSG00000117984","properties":[{"key":"ExpressionPatterns","value":"Low tissue specificity"}]},{"database":"MalaCards","id":"CTSD","properties":[{"key":"Description","value":"-"}]},{"database":"MIM","id":"116840","properties":[{"key":"Type","value":"gene"}]},{"database":"MIM","id":"610127","properties":[{"key":"Type","value":"phenotype"}]},{"database":"OpenTargets","id":"ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"Orphanet","id":"700487","properties":[{"key":"Disease","value":"Congenital CLN10 disease"}]},{"database":"Orphanet","id":"700497","properties":[{"key":"Disease","value":"Juvenile CLN10 disease"}]},{"database":"Orphanet","id":"700492","properties":[{"key":"Disease","value":"Late infantile CLN10 disease"}]},{"database":"VEuPathDB","id":"HostDB:ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"eggNOG","id":"KOG1339","properties":[{"key":"ToxonomicScope","value":"Eukaryota"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"HOGENOM","id":"CLU_013253_3_3_1","properties":[{"key":"Description","value":"-"}]},{"database":"InParanoid","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"OMA","id":"KYDHDAS","properties":[{"key":"Fingerprint","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"PAN-GO","id":"P07339","properties":[{"key":"Number of GO annotations","value":"4 GO annotations based on evolutionary models"}]},{"database":"PhylomeDB","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"BioCyc","id":"MetaCyc:HS04183-MONOMER","properties":[{"key":"Description","value":"-"}]},{"database":"BRENDA","id":"3.4.23.5","properties":[{"key":"OrganismId","value":"2681"}]},{"database":"PathwayCommons","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"Reactome","id":"R-HSA-1442490","properties":[{"key":"PathwayName","value":"Collagen degradation"}]},{"database":"Reactome","id":"R-HSA-2022377","properties":[{"key":"PathwayName","value":"Metabolism of Angiotensinogen to Angiotensins"}]},{"database":"Reactome","id":"R-HSA-2132295","properties":[{"key":"PathwayName","value":"MHC class II antigen presentation"}]},{"database":"Reactome","id":"R-HSA-6798695","properties":[{"key":"PathwayName","value":"Neutrophil degranulation"}]},{"database":"Reactome","id":"R-HSA-77387","properties":[{"key":"PathwayName","value":"Insulin receptor recycling"}]},{"database":"Reactome","id":"R-HSA-9018519","properties":[{"key":"PathwayName","value":"Estrogen-dependent gene expression"}]},{"database":"SignaLink","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"SIGNOR","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"Agora","id":"ENSG00000117984","properties":[{"key":"NominatedTarget","value":"-"}]},{"database":"BioGRID-ORCS","id":"1509","properties":[{"key":"hits","value":"9 hits in 1161 CRISPR screens"}]},{"database":"ChiTaRS","id":"CTSD","properties":[{"key":"OrganismName","value":"human"}]},{"database":"EvolutionaryTrace","id":"P07339","properties":[{"key":"Description","value":"-"}]},{"database":"GeneWiki","id":"Cathepsin_D","properties":[{"key":"Description","value":"-"}]},{"database":"GenomeRNAi","id":"1509","properties":[{"key":"Description","value":"-"}]},{"database":"Pharos","id":"P07339","properties":[{"key":"DevelopmentLevel","value":"Tchem"}]},{"database":"PRO","id":"PR:P07339","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"RNAct","id":"P07339","properties":[{"key":"moleculeType","value":"protein"}]},{"database":"Bgee","id":"ENSG00000117984","properties":[{"key":"ExpressionPatterns","value":"Expressed in right adrenal gland cortex and 199 other cell types or tissues"}]},{"database":"ExpressionAtlas","id":"P07339","properties":[{"key":"ExpressionPatterns","value":"baseline and differential"}]},{"database":"GO","id":"GO:0005829","properties":[{"key":"GoTerm","value":"C:cytosol"},{"key":"GoEvidenceType","value":"IDA:ARUK-UCL"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"12107093"}]},{"database":"GO","id":"GO:0031904","properties":[{"key":"GoTerm","value":"C:endosome lumen"},{"key":"GoEvidenceType","value":"IEA:Ensembl"}]},{"database":"GO","id":"GO:0010008","properties":[{"key":"GoTerm","value":"C:endosome membrane"},{"key":"GoEvidenceType","value":"IDA:ARUK-UCL"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"17188016"}]},{"database":"GO","id":"GO:0070062","properties":[{"key":"GoTerm","value":"C:extracellular exosome"},{"key":"GoEvidenceType","value":"HDA:UniProtKB"}],"evidences":[{"evidenceCode":"ECO:0007005","source":"PubMed","id":"19056867"},{"evidenceCode":"ECO:0007005","source":"PubMed","id":"23533145"}]},{"database":"GO","id":"GO:0031012","properties":[{"key":"GoTerm","value":"C:extracellular matrix"},{"key":"GoEvidenceType","value":"HDA:BHF-UCL"}],"evidences":[{"evidenceCode":"ECO:0007005","source":"PubMed","id":"25037231"},{"evidenceCode":"ECO:0007005","source":"PubMed","id":"28344315"},{"evidenceCode":"ECO:0007005","source":"PubMed","id":"28675934"}]},{"database":"GO","id":"GO:0005576","properties":[{"key":"GoTerm","value":"C:extracellular region"},{"key":"GoEvidenceType","value":"HDA:UniProtKB"}],"evidences":[{"evidenceCode":"ECO:0007005","source":"PubMed","id":"16502470"}]},{"database":"GO","id":"GO:1904813","properties":[{"key":"GoTerm","value":"C:ficolin-1-rich granule lumen"},{"key":"GoEvidenceType","value":"TAS:Reactome"}]},{"database":"GO","id":"GO:0043202","properties":[{"key":"GoTerm","value":"C:lysosomal lumen"},{"key":"GoEvidenceType","value":"TAS:Reactome"}]},{"database":"GO","id":"GO:0005765","properties":[{"key":"GoTerm","value":"C:lysosomal membrane"},{"key":"GoEvidenceType","value":"IDA:ARUK-UCL"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"17188016"}]},{"database":"GO","id":"GO:0005764","properties":[{"key":"GoTerm","value":"C:lysosome"},{"key":"GoEvidenceType","value":"IDA:UniProtKB"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"1837142"}]},{"database":"GO","id":"GO:0042470","properties":[{"key":"GoTerm","value":"C:melanosome"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-SubCell"}]},{"database":"GO","id":"GO:0045121","properties":[{"key":"GoTerm","value":"C:membrane raft"},{"key":"GoEvidenceType","value":"IDA:UniProtKB"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"25204797"}]},{"database":"GO","id":"GO:0035580","properties":[{"key":"GoTerm","value":"C:specific granule lumen"},{"key":"GoEvidenceType","value":"TAS:Reactome"}]},{"database":"GO","id":"GO:1904724","properties":[{"key":"GoTerm","value":"C:tertiary granule lumen"},{"key":"GoEvidenceType","value":"TAS:Reactome"}]},{"database":"GO","id":"GO:0004190","properties":[{"key":"GoTerm","value":"F:aspartic-type endopeptidase activity"},{"key":"GoEvidenceType","value":"IBA:GO_Central"}]},{"database":"GO","id":"GO:0070001","properties":[{"key":"GoTerm","value":"F:aspartic-type peptidase activity"},{"key":"GoEvidenceType","value":"ISS:ARUK-UCL"}]},{"database":"GO","id":"GO:0004197","properties":[{"key":"GoTerm","value":"F:cysteine-type endopeptidase activity"},{"key":"GoEvidenceType","value":"TAS:Reactome"}]},{"database":"GO","id":"GO:0008233","properties":[{"key":"GoTerm","value":"F:peptidase activity"},{"key":"GoEvidenceType","value":"IDA:ARUK-UCL"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"16997486"}]},{"database":"GO","id":"GO:0019886","properties":[{"key":"GoTerm","value":"P:antigen processing and presentation of exogenous peptide antigen via MHC class II"},{"key":"GoEvidenceType","value":"TAS:Reactome"}]},{"database":"GO","id":"GO:0097194","properties":[{"key":"GoTerm","value":"P:execution phase of apoptosis"},{"key":"GoEvidenceType","value":"IDA:ARUK-UCL"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"12107093"}]},{"database":"GO","id":"GO:1901143","properties":[{"key":"GoTerm","value":"P:insulin catabolic process"},{"key":"GoEvidenceType","value":"IEA:Ensembl"}]},{"database":"GO","id":"GO:0038020","properties":[{"key":"GoTerm","value":"P:insulin receptor recycling"},{"key":"GoEvidenceType","value":"IEA:Ensembl"}]},{"database":"GO","id":"GO:0042159","properties":[{"key":"GoTerm","value":"P:lipoprotein catabolic process"},{"key":"GoEvidenceType","value":"IDA:ARUK-UCL"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"16997486"}]},{"database":"GO","id":"GO:0043065","properties":[{"key":"GoTerm","value":"P:positive regulation of apoptotic process"},{"key":"GoEvidenceType","value":"IDA:ARUK-UCL"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"12107093"}]},{"database":"GO","id":"GO:0006508","properties":[{"key":"GoTerm","value":"P:proteolysis"},{"key":"GoEvidenceType","value":"IDA:ARUK-UCL"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"16997486"}]},{"database":"GO","id":"GO:0070201","properties":[{"key":"GoTerm","value":"P:regulation of establishment of protein localization"},{"key":"GoEvidenceType","value":"IDA:ARUK-UCL"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"12107093"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":412,"molWeight":44552,"crc64":"903FB8412E0CF0B0","md5":"96FAAB77E7A636425952954C519DFE4B"},"extraAttributes":{"countByCommentType":{"FUNCTION":1,"CATALYTIC ACTIVITY":1,"SUBUNIT":1,"INTERACTION":30,"SUBCELLULAR LOCATION":1,"TISSUE SPECIFICITY":1,"PTM":3,"POLYMORPHISM":1,"DISEASE":1,"SIMILARITY":1,"WEB RESOURCE":1},"countByFeatureType":{"Signal":1,"Propeptide":1,"Chain":3,"Domain":1,"Active site":2,"Glycosylation":3,"Disulfide bond":4,"Natural variant":4,"Beta strand":28,"Turn":6,"Helix":13},"uniParcId":"UPI00000463E2"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"C9JH19","uniProtkbId":"C9JH19_HUMAN","entryAudit":{"firstPublicDate":"2009-11-03","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2021-04-07","entryVersion":111,"sequenceVersion":2},"annotationScore":3.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000415036.2"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000415036.2"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":451,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5029878290"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":446,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Region","location":{"start":{"value":276,"modifier":"EXACT"},"end":{"value":302,"modifier":"EXACT"}},"description":"Disordered","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"MobiDB-lite"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":334,"modifier":"EXACT"},"end":{"value":334,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Glycosylation","location":{"start":{"value":134,"modifier":"EXACT"},"end":{"value":134,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Glycosylation","location":{"start":{"value":263,"modifier":"EXACT"},"end":{"value":263,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Disulfide bond","location":{"start":{"value":91,"modifier":"EXACT"},"end":{"value":160,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-2"}]},{"type":"Disulfide bond","location":{"start":{"value":110,"modifier":"EXACT"},"end":{"value":117,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":325,"modifier":"EXACT"},"end":{"value":329,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-2"}]},{"type":"Disulfide bond","location":{"start":{"value":368,"modifier":"EXACT"},"end":{"value":405,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"},{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"C9JH19"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"C9JH19"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022729"},{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000415036.2"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000415036.2"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000415036.2"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"21269460","citationType":"journal article","authors":["Burkard T.R.","Planyavsky M.","Kaupe I.","Breitwieser F.P.","Burckstummer T.","Bennett K.L.","Superti-Furga G.","Colinge J."],"citationCrossReferences":[{"database":"PubMed","id":"21269460"},{"database":"DOI","id":"10.1186/1752-0509-5-17"}],"title":"Initial characterization of the human central proteome.","publicationDate":"2011","journal":"BMC Syst. Biol.","firstPage":"17","lastPage":"17","volume":"5"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"21269460"}]},{"referenceNumber":5,"citation":{"id":"24275569","citationType":"journal article","authors":["Bian Y.","Song C.","Cheng K.","Dong M.","Wang F.","Huang J.","Sun D.","Wang L.","Ye M.","Zou H."],"citationCrossReferences":[{"database":"PubMed","id":"24275569"},{"database":"DOI","id":"10.1016/j.jprot.2013.11.014"}],"title":"An enzyme assisted RP-RPLC approach for in-depth analysis of human liver phosphoproteome.","publicationDate":"2014","journal":"J. Proteomics","firstPage":"253","lastPage":"262","volume":"96"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"24275569"}]},{"referenceNumber":6,"citation":{"id":"25944712","citationType":"journal article","authors":["Vaca Jacome A.S.","Rabilloud T.","Schaeffer-Reiss C.","Rompais M.","Ayoub D.","Lane L.","Bairoch A.","Van Dorsselaer A.","Carapito C."],"citationCrossReferences":[{"database":"PubMed","id":"25944712"}],"title":"N-terminome analysis of the human mitochondrial proteome.","publicationDate":"2015","journal":"Proteomics","firstPage":"2519","lastPage":"2524","volume":"15"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"25944712"}]},{"referenceNumber":7,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000415036.2"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"AlphaFoldDB","id":"C9JH19","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"C9JH19","properties":[{"key":"Description","value":"-"}]},{"database":"GlyCosmos","id":"C9JH19","properties":[{"key":"glycosylation","value":"2 sites, No reported glycans"}]},{"database":"MassIVE","id":"C9JH19","properties":[{"key":"Description","value":"-"}]},{"database":"ProteomicsDB","id":"10167","properties":[{"key":"Description","value":"-"}]},{"database":"Antibodypedia","id":"880","properties":[{"key":"antibodies","value":"1212 antibodies from 49 providers"}]},{"database":"Ensembl","id":"ENST00000438213.6","properties":[{"key":"ProteinId","value":"ENSP00000415036.2"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"UCSC","id":"uc057xtl.1","properties":[{"key":"OrganismName","value":"human"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"VEuPathDB","id":"HostDB:ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"ChiTaRS","id":"CTSD","properties":[{"key":"OrganismName","value":"human"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"Bgee","id":"ENSG00000117984","properties":[{"key":"ExpressionPatterns","value":"Expressed in right adrenal gland cortex and 199 other cell types or tissues"}]},{"database":"ExpressionAtlas","id":"C9JH19","properties":[{"key":"ExpressionPatterns","value":"baseline and differential"}]},{"database":"GO","id":"GO:0005764","properties":[{"key":"GoTerm","value":"C:lysosome"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-SubCell"}]},{"database":"GO","id":"GO:0004190","properties":[{"key":"GoTerm","value":"F:aspartic-type endopeptidase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-EC"}]},{"database":"GO","id":"GO:0006508","properties":[{"key":"GoTerm","value":"P:proteolysis"},{"key":"GoEvidenceType","value":"IEA:InterPro"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000066","properties":[{"key":"EntryName","value":"Napsin A aspartic peptidase"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"2"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQSPEGRRVQPSQGRGSQGGALGPQGTLQSRQGLHALTPRRVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":451,"molWeight":48614,"crc64":"0D4E976059A61229","md5":"F3B77CC51590EB5226E4DE50491D9096"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Region":1,"Active site":2,"Glycosylation":2,"Disulfide bond":4},"uniParcId":"UPI0018BB63E5"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"F8W787","secondaryAccessions":["H7C1V0"],"uniProtkbId":"F8W787_HUMAN","entryAudit":{"firstPublicDate":"2011-09-21","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2021-04-07","entryVersion":98,"sequenceVersion":2},"annotationScore":3.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000356164.4"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000356164.4"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Domain","location":{"start":{"value":44,"modifier":"EXACT"},"end":{"value":372,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":62,"modifier":"EXACT"},"end":{"value":62,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":260,"modifier":"EXACT"},"end":{"value":260,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Glycosylation","location":{"start":{"value":99,"modifier":"EXACT"},"end":{"value":99,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Glycosylation","location":{"start":{"value":228,"modifier":"EXACT"},"end":{"value":228,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Disulfide bond","location":{"start":{"value":56,"modifier":"EXACT"},"end":{"value":125,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-2"}]},{"type":"Disulfide bond","location":{"start":{"value":75,"modifier":"EXACT"},"end":{"value":82,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":251,"modifier":"EXACT"},"end":{"value":255,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":294,"modifier":"EXACT"},"end":{"value":331,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"},{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"F8W787"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"F8W787"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000356164.4"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000356164.4"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000356164.4"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"21269460","citationType":"journal article","authors":["Burkard T.R.","Planyavsky M.","Kaupe I.","Breitwieser F.P.","Burckstummer T.","Bennett K.L.","Superti-Furga G.","Colinge J."],"citationCrossReferences":[{"database":"PubMed","id":"21269460"},{"database":"DOI","id":"10.1186/1752-0509-5-17"}],"title":"Initial characterization of the human central proteome.","publicationDate":"2011","journal":"BMC Syst. Biol.","firstPage":"17","lastPage":"17","volume":"5"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"21269460"}]},{"referenceNumber":5,"citation":{"id":"24275569","citationType":"journal article","authors":["Bian Y.","Song C.","Cheng K.","Dong M.","Wang F.","Huang J.","Sun D.","Wang L.","Ye M.","Zou H."],"citationCrossReferences":[{"database":"PubMed","id":"24275569"},{"database":"DOI","id":"10.1016/j.jprot.2013.11.014"}],"title":"An enzyme assisted RP-RPLC approach for in-depth analysis of human liver phosphoproteome.","publicationDate":"2014","journal":"J. Proteomics","firstPage":"253","lastPage":"262","volume":"96"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"24275569"}]},{"referenceNumber":6,"citation":{"id":"25944712","citationType":"journal article","authors":["Vaca Jacome A.S.","Rabilloud T.","Schaeffer-Reiss C.","Rompais M.","Ayoub D.","Lane L.","Bairoch A.","Van Dorsselaer A.","Carapito C."],"citationCrossReferences":[{"database":"PubMed","id":"25944712"}],"title":"N-terminome analysis of the human mitochondrial proteome.","publicationDate":"2015","journal":"Proteomics","firstPage":"2519","lastPage":"2524","volume":"15"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"25944712"}]},{"referenceNumber":7,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000356164.4"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"AlphaFoldDB","id":"F8W787","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"F8W787","properties":[{"key":"Description","value":"-"}]},{"database":"ProteomicsDB","id":"29911","properties":[{"key":"Description","value":"-"}]},{"database":"ProteomicsDB","id":"44767","properties":[{"key":"Description","value":"-"}]},{"database":"Antibodypedia","id":"880","properties":[{"key":"antibodies","value":"1212 antibodies from 49 providers"}]},{"database":"Ensembl","id":"ENST00000367196.4","properties":[{"key":"ProteinId","value":"ENSP00000356164.4"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"Ensembl","id":"ENST00000429746.2","properties":[{"key":"ProteinId","value":"ENSP00000402586.2"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"UCSC","id":"uc057xtj.1","properties":[{"key":"OrganismName","value":"human"}]},{"database":"UCSC","id":"uc057xtm.1","properties":[{"key":"OrganismName","value":"human"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"OpenTargets","id":"ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"VEuPathDB","id":"HostDB:ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"HOGENOM","id":"CLU_013253_5_2_1","properties":[{"key":"Description","value":"-"}]},{"database":"HOGENOM","id":"CLU_013253_6_2_1","properties":[{"key":"Description","value":"-"}]},{"database":"OMA","id":"KYDHDAS","properties":[{"key":"Fingerprint","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"ChiTaRS","id":"CTSD","properties":[{"key":"OrganismName","value":"human"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"Bgee","id":"ENSG00000117984","properties":[{"key":"ExpressionPatterns","value":"Expressed in right adrenal gland cortex and 199 other cell types or tissues"}]},{"database":"GO","id":"GO:0005764","properties":[{"key":"GoTerm","value":"C:lysosome"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-SubCell"}]},{"database":"GO","id":"GO:0004190","properties":[{"key":"GoTerm","value":"F:aspartic-type endopeptidase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-EC"}]},{"database":"GO","id":"GO:0006508","properties":[{"key":"GoTerm","value":"P:proteolysis"},{"key":"GoEvidenceType","value":"IEA:InterPro"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.60.40.1960:FF:000001","properties":[{"key":"EntryName","value":"Cathepsin D"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":377,"molWeight":40784,"crc64":"C57554E98ADB2E57","md5":"306C1A3C8BDDBE02BE8FC0198BD5502E"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Domain":1,"Active site":2,"Glycosylation":2,"Disulfide bond":4},"uniParcId":"UPI00001F9A9C"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"V9HWI3","uniProtkbId":"V9HWI3_HUMAN","entryAudit":{"firstPublicDate":"2014-03-19","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2014-03-19","entryVersion":92,"sequenceVersion":1},"annotationScore":3.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"ACJ13652.1"}],"lineage":["Eukaryota","Metazoa","Chordata","Craniata","Vertebrata","Euteleostomi","Mammalia","Eutheria","Euarchontoglires","Primates","Haplorrhini","Catarrhini","Hominidae","Homo"]},"proteinExistence":"2: Evidence at transcript level","proteinDescription":{"recommendedName":{"fullName":{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"ACJ13652.1"}],"value":"HEL-S-130P"},"synonyms":[{"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"ALQ33465.1"}],"value":"CTSD"}],"orfNames":[{"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"EAX02461.1"}],"value":"hCG_49857"}]}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":412,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5014314732"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":407,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":295,"modifier":"EXACT"},"end":{"value":295,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Glycosylation","location":{"start":{"value":134,"modifier":"EXACT"},"end":{"value":134,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Glycosylation","location":{"start":{"value":263,"modifier":"EXACT"},"end":{"value":263,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Disulfide bond","location":{"start":{"value":91,"modifier":"EXACT"},"end":{"value":160,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-2"}]},{"type":"Disulfide bond","location":{"start":{"value":110,"modifier":"EXACT"},"end":{"value":117,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":286,"modifier":"EXACT"},"end":{"value":290,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":329,"modifier":"EXACT"},"end":{"value":366,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"},{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11181995","citationType":"journal article","authors":["Venter J.C.","Adams M.D.","Myers E.W.","Li P.W.","Mural R.J.","Sutton G.G.","Smith H.O.","Yandell M.","Evans C.A.","Holt R.A.","Gocayne J.D.","Amanatides P.","Ballew R.M.","Huson D.H.","Wortman J.R.","Zhang Q.","Kodira C.D.","Zheng X.H.","Chen L.","Skupski M.","Subramanian G.","Thomas P.D.","Zhang J.","Gabor Miklos G.L.","Nelson C.","Broder S.","Clark A.G.","Nadeau J.","McKusick V.A.","Zinder N.","Levine A.J.","Roberts R.J.","Simon M.","Slayman C.","Hunkapiller M.","Bolanos R.","Delcher A.","Dew I.","Fasulo D.","Flanigan M.","Florea L.","Halpern A.","Hannenhalli S.","Kravitz S.","Levy S.","Mobarry C.","Reinert K.","Remington K.","Abu-Threideh J.","Beasley E.","Biddick K.","Bonazzi V.","Brandon R.","Cargill M.","Chandramouliswaran I.","Charlab R.","Chaturvedi K.","Deng Z.","Di Francesco V.","Dunn P.","Eilbeck K.","Evangelista C.","Gabrielian A.E.","Gan W.","Ge W.","Gong F.","Gu Z.","Guan P.","Heiman T.J.","Higgins M.E.","Ji R.R.","Ke Z.","Ketchum K.A.","Lai Z.","Lei Y.","Li Z.","Li J.","Liang Y.","Lin X.","Lu F.","Merkulov G.V.","Milshina N.","Moore H.M.","Naik A.K.","Narayan V.A.","Neelam B.","Nusskern D.","Rusch D.B.","Salzberg S.","Shao W.","Shue B.","Sun J.","Wang Z.","Wang A.","Wang X.","Wang J.","Wei M.","Wides R.","Xiao C.","Yan C.","Yao A.","Ye J.","Zhan M.","Zhang W.","Zhang H.","Zhao Q.","Zheng L.","Zhong F.","Zhong W.","Zhu S.","Zhao S.","Gilbert D.","Baumhueter S.","Spier G.","Carter C.","Cravchik A.","Woodage T.","Ali F.","An H.","Awe A.","Baldwin D.","Baden H.","Barnstead M.","Barrow I.","Beeson K.","Busam D.","Carver A.","Center A.","Cheng M.L.","Curry L.","Danaher S.","Davenport L.","Desilets R.","Dietz S.","Dodson K.","Doup L.","Ferriera S.","Garg N.","Gluecksmann A.","Hart B.","Haynes J.","Haynes C.","Heiner C.","Hladun S.","Hostin D.","Houck J.","Howland T.","Ibegwam C.","Johnson J.","Kalush F.","Kline L.","Koduru S.","Love A.","Mann F.","May D.","McCawley S.","McIntosh T.","McMullen I.","Moy M.","Moy L.","Murphy B.","Nelson K.","Pfannkoch C.","Pratts E.","Puri V.","Qureshi H.","Reardon M.","Rodriguez R.","Rogers Y.H.","Romblad D.","Ruhfel B.","Scott R.","Sitter C.","Smallwood M.","Stewart E.","Strong R.","Suh E.","Thomas R.","Tint N.N.","Tse S.","Vech C.","Wang G.","Wetter J.","Williams S.","Williams M.","Windsor S.","Winn-Deen E.","Wolfe K.","Zaveri J.","Zaveri K.","Abril J.F.","Guigo R.","Campbell M.J.","Sjolander K.V.","Karlak B.","Kejariwal A.","Mi H.","Lazareva B.","Hatton T.","Narechania A.","Diemer K.","Muruganujan A.","Guo N.","Sato S.","Bafna V.","Istrail S.","Lippert R.","Schwartz R.","Walenz B.","Yooseph S.","Allen D.","Basu A.","Baxendale J.","Blick L.","Caminha M.","Carnes-Stine J.","Caulk P.","Chiang Y.H.","Coyne M.","Dahlke C.","Mays A.","Dombroski M.","Donnelly M.","Ely D.","Esparham S.","Fosler C.","Gire H.","Glanowski S.","Glasser K.","Glodek A.","Gorokhov M.","Graham K.","Gropman B.","Harris M.","Heil J.","Henderson S.","Hoover J.","Jennings D.","Jordan C.","Jordan J.","Kasha J.","Kagan L.","Kraft C.","Levitsky A.","Lewis M.","Liu X.","Lopez J.","Ma D.","Majoros W.","McDaniel J.","Murphy S.","Newman M.","Nguyen T.","Nguyen N.","Nodell M.","Pan S.","Peck J.","Peterson M.","Rowe W.","Sanders R.","Scott J.","Simpson M.","Smith T.","Sprague A.","Stockwell T.","Turner R.","Venter E.","Wang M.","Wen M.","Wu D.","Wu M.","Xia A.","Zandieh A.","Zhu X."],"citationCrossReferences":[{"database":"PubMed","id":"11181995"},{"database":"DOI","id":"10.1126/science.1058040"}],"title":"The sequence of the human genome.","publicationDate":"2001","journal":"Science","firstPage":"1304","lastPage":"1351","volume":"291"},"referencePositions":["NUCLEOTIDE SEQUENCE"],"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"EAX02461.1"}]},{"referenceNumber":2,"citation":{"id":"CI-56FT96O07V0QG","citationType":"submission","authors":["Mural R.J.","Istrail S.","Sutton G.","Florea L.","Halpern A.L.","Mobarry C.M.","Lippert R.","Walenz B.","Shatkay H.","Dew I.","Miller J.R.","Flanigan M.J.","Edwards N.J.","Bolanos R.","Fasulo D.","Halldorsson B.V.","Hannenhalli S.","Turner R.","Yooseph S.","Lu F.","Nusskern D.R.","Shue B.C.","Zheng X.H.","Zhong F.","Delcher A.L.","Huson D.H.","Kravitz S.A.","Mouchard L.","Reinert K.","Remington K.A.","Clark A.G.","Waterman M.S.","Eichler E.E.","Adams M.D.","Hunkapiller M.W.","Myers E.W.","Venter J.C."],"publicationDate":"JUL-2005","submissionDatabase":"EMBL/GenBank/DDBJ databases"},"referencePositions":["NUCLEOTIDE SEQUENCE"],"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"EAX02461.1"}]},{"referenceNumber":3,"citation":{"id":"CI-FEI80TKHQ30BS","citationType":"submission","authors":["Li J.Y.","Wang H.Y.","Liu F.J.","Liu J."],"publicationDate":"JUN-2008","submissionDatabase":"EMBL/GenBank/DDBJ databases"},"referencePositions":["NUCLEOTIDE SEQUENCE"],"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"ACJ13652.1"}]},{"referenceNumber":4,"citation":{"id":"26871637","citationType":"journal article","authors":["Yang X.","Coulombe-Huntington J.","Kang S.","Sheynkman G.M.","Hao T.","Richardson A.","Sun S.","Yang F.","Shen Y.A.","Murray R.","Spirohn K.","Begg B.E.","Duran-Frigola M.","MacWilliams A.","Pevzner S.J.","Zhong Q.","Trigg S.A.","Tam S.","Ghamsari L.","Sahni N.","Yi S.","Rodriguez M.D.","Balcha D.","Tan G.","Costanzo M.","Andrews B.","Boone C.","Zhou X.J.","Salehi-Ashtiani K.","Charloteaux B.","Chen A.","Calderwood M.A.","Aloy P.","Roth F.P.","Hill D.E.","Iakoucheva L.M.","Xia Y.","Vidal M."],"citationCrossReferences":[{"database":"PubMed","id":"26871637"},{"database":"DOI","id":"10.1016/j.cell.2016.01.029"}],"title":"Widespread Expansion of Protein Interaction Capabilities by Alternative Splicing.","publicationDate":"2016","journal":"Cell","firstPage":"805","lastPage":"817","volume":"164"},"referencePositions":["NUCLEOTIDE SEQUENCE"],"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"ALQ33465.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"EU794598","properties":[{"key":"ProteinId","value":"ACJ13652.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"mRNA"}]},{"database":"EMBL","id":"KU178007","properties":[{"key":"ProteinId","value":"ALQ33465.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"mRNA"}]},{"database":"EMBL","id":"CH471158","properties":[{"key":"ProteinId","value":"EAX02461.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"RefSeq","id":"NP_001900.1","properties":[{"key":"NucleotideSequenceId","value":"NM_001909.5"}]},{"database":"AlphaFoldDB","id":"V9HWI3","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"V9HWI3","properties":[{"key":"Description","value":"-"}]},{"database":"IntAct","id":"V9HWI3","properties":[{"key":"Interactions","value":"2"}]},{"database":"MEROPS","id":"A01.009","properties":[{"key":"Description","value":"-"}]},{"database":"Antibodypedia","id":"880","properties":[{"key":"antibodies","value":"1212 antibodies from 49 providers"}]},{"database":"DNASU","id":"1509","properties":[{"key":"Description","value":"-"}]},{"database":"GeneID","id":"1509","properties":[{"key":"Description","value":"-"}]},{"database":"KEGG","id":"hsa:1509","properties":[{"key":"Description","value":"-"}]},{"database":"UCSC","id":"uc001luc.2","properties":[{"key":"OrganismName","value":"human"}]},{"database":"CTD","id":"1509","properties":[{"key":"Description","value":"-"}]},{"database":"DisGeNET","id":"1509","properties":[{"key":"Description","value":"-"}]},{"database":"VEuPathDB","id":"HostDB:ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"HOGENOM","id":"CLU_013253_3_3_1","properties":[{"key":"Description","value":"-"}]},{"database":"OMA","id":"KYDHDAS","properties":[{"key":"Fingerprint","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"BioGRID-ORCS","id":"1509","properties":[{"key":"hits","value":"9 hits in 1161 CRISPR screens"}]},{"database":"ChiTaRS","id":"CTSD","properties":[{"key":"OrganismName","value":"human"}]},{"database":"ExpressionAtlas","id":"V9HWI3","properties":[{"key":"ExpressionPatterns","value":"baseline and differential"}]},{"database":"GO","id":"GO:0031904","properties":[{"key":"GoTerm","value":"C:endosome lumen"},{"key":"GoEvidenceType","value":"IEA:Ensembl"}]},{"database":"GO","id":"GO:0005576","properties":[{"key":"GoTerm","value":"C:extracellular region"},{"key":"GoEvidenceType","value":"IEA:Ensembl"}]},{"database":"GO","id":"GO:0005764","properties":[{"key":"GoTerm","value":"C:lysosome"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-SubCell"}]},{"database":"GO","id":"GO:0004190","properties":[{"key":"GoTerm","value":"F:aspartic-type endopeptidase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-EC"}]},{"database":"GO","id":"GO:1901143","properties":[{"key":"GoTerm","value":"P:insulin catabolic process"},{"key":"GoEvidenceType","value":"IEA:Ensembl"}]},{"database":"GO","id":"GO:0038020","properties":[{"key":"GoTerm","value":"P:insulin receptor recycling"},{"key":"GoEvidenceType","value":"IEA:Ensembl"}]},{"database":"GO","id":"GO:0006508","properties":[{"key":"GoTerm","value":"P:proteolysis"},{"key":"GoEvidenceType","value":"IEA:InterPro"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":412,"molWeight":44552,"crc64":"903FB8412E0CF0B0","md5":"96FAAB77E7A636425952954C519DFE4B"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2,"Glycosylation":2,"Disulfide bond":4},"uniParcId":"UPI00000463E2"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0A1B0GW44","uniProtkbId":"A0A1B0GW44_HUMAN","entryAudit":{"firstPublicDate":"2016-10-05","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2016-10-05","entryVersion":51,"sequenceVersion":1},"annotationScore":3.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490770.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490770.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":405,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5008408761"},{"type":"Domain","location":{"start":{"value":72,"modifier":"EXACT"},"end":{"value":400,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":90,"modifier":"EXACT"},"end":{"value":90,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":288,"modifier":"EXACT"},"end":{"value":288,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Glycosylation","location":{"start":{"value":127,"modifier":"EXACT"},"end":{"value":127,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Glycosylation","location":{"start":{"value":256,"modifier":"EXACT"},"end":{"value":256,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Disulfide bond","location":{"start":{"value":84,"modifier":"EXACT"},"end":{"value":153,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-2"}]},{"type":"Disulfide bond","location":{"start":{"value":103,"modifier":"EXACT"},"end":{"value":110,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":279,"modifier":"EXACT"},"end":{"value":283,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":322,"modifier":"EXACT"},"end":{"value":359,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"},{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0A1B0GW44"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0A1B0GW44"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490770.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490770.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490770.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"21269460","citationType":"journal article","authors":["Burkard T.R.","Planyavsky M.","Kaupe I.","Breitwieser F.P.","Burckstummer T.","Bennett K.L.","Superti-Furga G.","Colinge J."],"citationCrossReferences":[{"database":"PubMed","id":"21269460"},{"database":"DOI","id":"10.1186/1752-0509-5-17"}],"title":"Initial characterization of the human central proteome.","publicationDate":"2011","journal":"BMC Syst. Biol.","firstPage":"17","lastPage":"17","volume":"5"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"21269460"}]},{"referenceNumber":5,"citation":{"id":"25944712","citationType":"journal article","authors":["Vaca Jacome A.S.","Rabilloud T.","Schaeffer-Reiss C.","Rompais M.","Ayoub D.","Lane L.","Bairoch A.","Van Dorsselaer A.","Carapito C."],"citationCrossReferences":[{"database":"PubMed","id":"25944712"}],"title":"N-terminome analysis of the human mitochondrial proteome.","publicationDate":"2015","journal":"Proteomics","firstPage":"2519","lastPage":"2524","volume":"15"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"25944712"}]},{"referenceNumber":6,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490770.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"AlphaFoldDB","id":"A0A1B0GW44","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"A0A1B0GW44","properties":[{"key":"Description","value":"-"}]},{"database":"GlyCosmos","id":"A0A1B0GW44","properties":[{"key":"glycosylation","value":"2 sites, No reported glycans"}]},{"database":"MassIVE","id":"A0A1B0GW44","properties":[{"key":"Description","value":"-"}]},{"database":"Antibodypedia","id":"880","properties":[{"key":"antibodies","value":"1212 antibodies from 49 providers"}]},{"database":"Ensembl","id":"ENST00000636571.1","properties":[{"key":"ProteinId","value":"ENSP00000490770.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"VEuPathDB","id":"HostDB:ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"ChiTaRS","id":"CTSD","properties":[{"key":"OrganismName","value":"human"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"Bgee","id":"ENSG00000117984","properties":[{"key":"ExpressionPatterns","value":"Expressed in right adrenal gland cortex and 199 other cell types or tissues"}]},{"database":"ExpressionAtlas","id":"A0A1B0GW44","properties":[{"key":"ExpressionPatterns","value":"baseline and differential"}]},{"database":"GO","id":"GO:0005764","properties":[{"key":"GoTerm","value":"C:lysosome"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-SubCell"}]},{"database":"GO","id":"GO:0004190","properties":[{"key":"GoTerm","value":"F:aspartic-type endopeptidase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-EC"}]},{"database":"GO","id":"GO:0006508","properties":[{"key":"GoTerm","value":"P:proteolysis"},{"key":"GoEvidenceType","value":"IEA:InterPro"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":405,"molWeight":43688,"crc64":"5C06A6AE9C48AD79","md5":"8A947DE86343FFC9287F698FBC62A57F"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2,"Glycosylation":2,"Disulfide bond":4},"uniParcId":"UPI0007E52BE2"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"F8WD96","uniProtkbId":"F8WD96_HUMAN","entryAudit":{"firstPublicDate":"2011-09-21","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2011-09-21","entryVersion":91,"sequenceVersion":1},"annotationScore":3.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000404902.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000404902.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":276,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5003379681"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":276,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Disulfide bond","location":{"start":{"value":110,"modifier":"EXACT"},"end":{"value":117,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"},{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"F8WD96"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"F8WD96"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000404902.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000404902.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000404902.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"21269460","citationType":"journal article","authors":["Burkard T.R.","Planyavsky M.","Kaupe I.","Breitwieser F.P.","Burckstummer T.","Bennett K.L.","Superti-Furga G.","Colinge J."],"citationCrossReferences":[{"database":"PubMed","id":"21269460"},{"database":"DOI","id":"10.1186/1752-0509-5-17"}],"title":"Initial characterization of the human central proteome.","publicationDate":"2011","journal":"BMC Syst. Biol.","firstPage":"17","lastPage":"17","volume":"5"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"21269460"}]},{"referenceNumber":5,"citation":{"id":"24275569","citationType":"journal article","authors":["Bian Y.","Song C.","Cheng K.","Dong M.","Wang F.","Huang J.","Sun D.","Wang L.","Ye M.","Zou H."],"citationCrossReferences":[{"database":"PubMed","id":"24275569"},{"database":"DOI","id":"10.1016/j.jprot.2013.11.014"}],"title":"An enzyme assisted RP-RPLC approach for in-depth analysis of human liver phosphoproteome.","publicationDate":"2014","journal":"J. Proteomics","firstPage":"253","lastPage":"262","volume":"96"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"24275569"}]},{"referenceNumber":6,"citation":{"id":"25944712","citationType":"journal article","authors":["Vaca Jacome A.S.","Rabilloud T.","Schaeffer-Reiss C.","Rompais M.","Ayoub D.","Lane L.","Bairoch A.","Van Dorsselaer A.","Carapito C."],"citationCrossReferences":[{"database":"PubMed","id":"25944712"}],"title":"N-terminome analysis of the human mitochondrial proteome.","publicationDate":"2015","journal":"Proteomics","firstPage":"2519","lastPage":"2524","volume":"15"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"25944712"}]},{"referenceNumber":7,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000404902.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"AlphaFoldDB","id":"F8WD96","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"F8WD96","properties":[{"key":"Description","value":"-"}]},{"database":"MassIVE","id":"F8WD96","properties":[{"key":"Description","value":"-"}]},{"database":"ProteomicsDB","id":"31420","properties":[{"key":"Description","value":"-"}]},{"database":"Ensembl","id":"ENST00000433655.6","properties":[{"key":"ProteinId","value":"ENSP00000404902.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"UCSC","id":"uc057xti.1","properties":[{"key":"OrganismName","value":"human"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"VEuPathDB","id":"HostDB:ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"OMA","id":"DIACCEL","properties":[{"key":"Fingerprint","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"ChiTaRS","id":"CTSD","properties":[{"key":"OrganismName","value":"human"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"Bgee","id":"ENSG00000117984","properties":[{"key":"ExpressionPatterns","value":"Expressed in right adrenal gland cortex and 199 other cell types or tissues"}]},{"database":"ExpressionAtlas","id":"F8WD96","properties":[{"key":"ExpressionPatterns","value":"baseline and differential"}]},{"database":"GO","id":"GO:0005764","properties":[{"key":"GoTerm","value":"C:lysosome"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-SubCell"}]},{"database":"GO","id":"GO:0004190","properties":[{"key":"GoTerm","value":"F:aspartic-type endopeptidase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-EC"}]},{"database":"GO","id":"GO:0006508","properties":[{"key":"GoTerm","value":"P:proteolysis"},{"key":"GoEvidenceType","value":"IEA:InterPro"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQ","length":276,"molWeight":30000,"crc64":"C78C9E359BB3C92D","md5":"6E8675AED46B12FF47D7DCEB07B4A6FE"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":1,"Disulfide bond":1},"uniParcId":"UPI0000D4DE30"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0A1B0GVD5","uniProtkbId":"A0A1B0GVD5_HUMAN","entryAudit":{"firstPublicDate":"2016-10-05","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2016-10-05","entryVersion":52,"sequenceVersion":1},"annotationScore":3.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490471.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490471.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":409,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5008408753"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":404,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":295,"modifier":"EXACT"},"end":{"value":295,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Glycosylation","location":{"start":{"value":134,"modifier":"EXACT"},"end":{"value":134,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Glycosylation","location":{"start":{"value":263,"modifier":"EXACT"},"end":{"value":263,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Disulfide bond","location":{"start":{"value":91,"modifier":"EXACT"},"end":{"value":160,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-2"}]},{"type":"Disulfide bond","location":{"start":{"value":110,"modifier":"EXACT"},"end":{"value":117,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":286,"modifier":"EXACT"},"end":{"value":290,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":329,"modifier":"EXACT"},"end":{"value":363,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"},{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0A1B0GVD5"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0A1B0GVD5"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490471.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490471.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490471.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"21269460","citationType":"journal article","authors":["Burkard T.R.","Planyavsky M.","Kaupe I.","Breitwieser F.P.","Burckstummer T.","Bennett K.L.","Superti-Furga G.","Colinge J."],"citationCrossReferences":[{"database":"PubMed","id":"21269460"},{"database":"DOI","id":"10.1186/1752-0509-5-17"}],"title":"Initial characterization of the human central proteome.","publicationDate":"2011","journal":"BMC Syst. Biol.","firstPage":"17","lastPage":"17","volume":"5"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"21269460"}]},{"referenceNumber":5,"citation":{"id":"25944712","citationType":"journal article","authors":["Vaca Jacome A.S.","Rabilloud T.","Schaeffer-Reiss C.","Rompais M.","Ayoub D.","Lane L.","Bairoch A.","Van Dorsselaer A.","Carapito C."],"citationCrossReferences":[{"database":"PubMed","id":"25944712"}],"title":"N-terminome analysis of the human mitochondrial proteome.","publicationDate":"2015","journal":"Proteomics","firstPage":"2519","lastPage":"2524","volume":"15"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"25944712"}]},{"referenceNumber":6,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490471.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"AlphaFoldDB","id":"A0A1B0GVD5","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"A0A1B0GVD5","properties":[{"key":"Description","value":"-"}]},{"database":"GlyCosmos","id":"A0A1B0GVD5","properties":[{"key":"glycosylation","value":"2 sites, No reported glycans"}]},{"database":"MassIVE","id":"A0A1B0GVD5","properties":[{"key":"Description","value":"-"}]},{"database":"Antibodypedia","id":"880","properties":[{"key":"antibodies","value":"1212 antibodies from 49 providers"}]},{"database":"Ensembl","id":"ENST00000637915.1","properties":[{"key":"ProteinId","value":"ENSP00000490471.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"VEuPathDB","id":"HostDB:ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"ChiTaRS","id":"CTSD","properties":[{"key":"OrganismName","value":"human"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"Bgee","id":"ENSG00000117984","properties":[{"key":"ExpressionPatterns","value":"Expressed in right adrenal gland cortex and 199 other cell types or tissues"}]},{"database":"ExpressionAtlas","id":"A0A1B0GVD5","properties":[{"key":"ExpressionPatterns","value":"baseline and differential"}]},{"database":"GO","id":"GO:0005764","properties":[{"key":"GoTerm","value":"C:lysosome"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-SubCell"}]},{"database":"GO","id":"GO:0004190","properties":[{"key":"GoTerm","value":"F:aspartic-type endopeptidase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-EC"}]},{"database":"GO","id":"GO:0006508","properties":[{"key":"GoTerm","value":"P:proteolysis"},{"key":"GoEvidenceType","value":"IEA:InterPro"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":409,"molWeight":44238,"crc64":"36DCD7D3ED13BD4E","md5":"BE883AEE44C02A6C3876270457935C35"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2,"Glycosylation":2,"Disulfide bond":4},"uniParcId":"UPI0007E52CCC"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0A1B0GVP3","uniProtkbId":"A0A1B0GVP3_HUMAN","entryAudit":{"firstPublicDate":"2016-10-05","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2016-10-05","entryVersion":51,"sequenceVersion":1},"annotationScore":3.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490598.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490598.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":405,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5008408757"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":400,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":295,"modifier":"EXACT"},"end":{"value":295,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Glycosylation","location":{"start":{"value":134,"modifier":"EXACT"},"end":{"value":134,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Glycosylation","location":{"start":{"value":263,"modifier":"EXACT"},"end":{"value":263,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Disulfide bond","location":{"start":{"value":91,"modifier":"EXACT"},"end":{"value":160,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-2"}]},{"type":"Disulfide bond","location":{"start":{"value":110,"modifier":"EXACT"},"end":{"value":117,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":286,"modifier":"EXACT"},"end":{"value":290,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"},{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0A1B0GVP3"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0A1B0GVP3"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490598.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490598.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490598.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"21269460","citationType":"journal article","authors":["Burkard T.R.","Planyavsky M.","Kaupe I.","Breitwieser F.P.","Burckstummer T.","Bennett K.L.","Superti-Furga G.","Colinge J."],"citationCrossReferences":[{"database":"PubMed","id":"21269460"},{"database":"DOI","id":"10.1186/1752-0509-5-17"}],"title":"Initial characterization of the human central proteome.","publicationDate":"2011","journal":"BMC Syst. Biol.","firstPage":"17","lastPage":"17","volume":"5"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"21269460"}]},{"referenceNumber":5,"citation":{"id":"25944712","citationType":"journal article","authors":["Vaca Jacome A.S.","Rabilloud T.","Schaeffer-Reiss C.","Rompais M.","Ayoub D.","Lane L.","Bairoch A.","Van Dorsselaer A.","Carapito C."],"citationCrossReferences":[{"database":"PubMed","id":"25944712"}],"title":"N-terminome analysis of the human mitochondrial proteome.","publicationDate":"2015","journal":"Proteomics","firstPage":"2519","lastPage":"2524","volume":"15"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"25944712"}]},{"referenceNumber":6,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490598.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"AlphaFoldDB","id":"A0A1B0GVP3","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"A0A1B0GVP3","properties":[{"key":"Description","value":"-"}]},{"database":"IntAct","id":"A0A1B0GVP3","properties":[{"key":"Interactions","value":"1"}]},{"database":"GlyCosmos","id":"A0A1B0GVP3","properties":[{"key":"glycosylation","value":"2 sites, No reported glycans"}]},{"database":"MassIVE","id":"A0A1B0GVP3","properties":[{"key":"Description","value":"-"}]},{"database":"Antibodypedia","id":"880","properties":[{"key":"antibodies","value":"1212 antibodies from 49 providers"}]},{"database":"Ensembl","id":"ENST00000637387.1","properties":[{"key":"ProteinId","value":"ENSP00000490598.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"VEuPathDB","id":"HostDB:ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"ChiTaRS","id":"CTSD","properties":[{"key":"OrganismName","value":"human"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"Bgee","id":"ENSG00000117984","properties":[{"key":"ExpressionPatterns","value":"Expressed in right adrenal gland cortex and 199 other cell types or tissues"}]},{"database":"ExpressionAtlas","id":"A0A1B0GVP3","properties":[{"key":"ExpressionPatterns","value":"baseline and differential"}]},{"database":"GO","id":"GO:0005764","properties":[{"key":"GoTerm","value":"C:lysosome"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-SubCell"}]},{"database":"GO","id":"GO:0004190","properties":[{"key":"GoTerm","value":"F:aspartic-type endopeptidase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-EC"}]},{"database":"GO","id":"GO:0006508","properties":[{"key":"GoTerm","value":"P:proteolysis"},{"key":"GoEvidenceType","value":"IEA:InterPro"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":405,"molWeight":43687,"crc64":"FAA490F788322EB9","md5":"CCFD9E329B79467CA5E042EA62399208"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2,"Glycosylation":2,"Disulfide bond":3},"uniParcId":"UPI0007E52D08"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0A1B0GWE8","uniProtkbId":"A0A1B0GWE8_HUMAN","entryAudit":{"firstPublicDate":"2016-10-05","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2016-10-05","entryVersion":51,"sequenceVersion":1},"annotationScore":3.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490897.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490897.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":410,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5008408691"},{"type":"Domain","location":{"start":{"value":77,"modifier":"EXACT"},"end":{"value":405,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":95,"modifier":"EXACT"},"end":{"value":95,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":293,"modifier":"EXACT"},"end":{"value":293,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Glycosylation","location":{"start":{"value":132,"modifier":"EXACT"},"end":{"value":132,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Glycosylation","location":{"start":{"value":261,"modifier":"EXACT"},"end":{"value":261,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Disulfide bond","location":{"start":{"value":89,"modifier":"EXACT"},"end":{"value":158,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-2"}]},{"type":"Disulfide bond","location":{"start":{"value":108,"modifier":"EXACT"},"end":{"value":115,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":284,"modifier":"EXACT"},"end":{"value":288,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":327,"modifier":"EXACT"},"end":{"value":364,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"},{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0A1B0GWE8"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0A1B0GWE8"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490897.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490897.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490897.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"21269460","citationType":"journal article","authors":["Burkard T.R.","Planyavsky M.","Kaupe I.","Breitwieser F.P.","Burckstummer T.","Bennett K.L.","Superti-Furga G.","Colinge J."],"citationCrossReferences":[{"database":"PubMed","id":"21269460"},{"database":"DOI","id":"10.1186/1752-0509-5-17"}],"title":"Initial characterization of the human central proteome.","publicationDate":"2011","journal":"BMC Syst. Biol.","firstPage":"17","lastPage":"17","volume":"5"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"21269460"}]},{"referenceNumber":5,"citation":{"id":"25944712","citationType":"journal article","authors":["Vaca Jacome A.S.","Rabilloud T.","Schaeffer-Reiss C.","Rompais M.","Ayoub D.","Lane L.","Bairoch A.","Van Dorsselaer A.","Carapito C."],"citationCrossReferences":[{"database":"PubMed","id":"25944712"}],"title":"N-terminome analysis of the human mitochondrial proteome.","publicationDate":"2015","journal":"Proteomics","firstPage":"2519","lastPage":"2524","volume":"15"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"25944712"}]},{"referenceNumber":6,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490897.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"AlphaFoldDB","id":"A0A1B0GWE8","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"A0A1B0GWE8","properties":[{"key":"Description","value":"-"}]},{"database":"GlyCosmos","id":"A0A1B0GWE8","properties":[{"key":"glycosylation","value":"2 sites, No reported glycans"}]},{"database":"MassIVE","id":"A0A1B0GWE8","properties":[{"key":"Description","value":"-"}]},{"database":"Antibodypedia","id":"880","properties":[{"key":"antibodies","value":"1212 antibodies from 49 providers"}]},{"database":"Ensembl","id":"ENST00000636843.1","properties":[{"key":"ProteinId","value":"ENSP00000490897.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"VEuPathDB","id":"HostDB:ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"ChiTaRS","id":"CTSD","properties":[{"key":"OrganismName","value":"human"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"Bgee","id":"ENSG00000117984","properties":[{"key":"ExpressionPatterns","value":"Expressed in right adrenal gland cortex and 199 other cell types or tissues"}]},{"database":"ExpressionAtlas","id":"A0A1B0GWE8","properties":[{"key":"ExpressionPatterns","value":"baseline and differential"}]},{"database":"GO","id":"GO:0005764","properties":[{"key":"GoTerm","value":"C:lysosome"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-SubCell"}]},{"database":"GO","id":"GO:0004190","properties":[{"key":"GoTerm","value":"F:aspartic-type endopeptidase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-EC"}]},{"database":"GO","id":"GO:0006508","properties":[{"key":"GoTerm","value":"P:proteolysis"},{"key":"GoEvidenceType","value":"IEA:InterPro"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":410,"molWeight":44353,"crc64":"15528FF44B00EDFC","md5":"26246B2206DB4128FABEA2FD35C4AED8"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2,"Glycosylation":2,"Disulfide bond":4},"uniParcId":"UPI0007E52AA4"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0A1B0GV23","uniProtkbId":"A0A1B0GV23_HUMAN","entryAudit":{"firstPublicDate":"2016-10-05","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2016-10-05","entryVersion":51,"sequenceVersion":1},"annotationScore":3.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490344.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490344.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":406,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5008408647"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":401,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":289,"modifier":"EXACT"},"end":{"value":289,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-1"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Glycosylation","location":{"start":{"value":134,"modifier":"EXACT"},"end":{"value":134,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Glycosylation","location":{"start":{"value":263,"modifier":"EXACT"},"end":{"value":263,"modifier":"EXACT"}},"description":"N-linked (GlcNAc...) asparagine","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-3"}],"featureId":""},{"type":"Disulfide bond","location":{"start":{"value":91,"modifier":"EXACT"},"end":{"value":160,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-2"}]},{"type":"Disulfide bond","location":{"start":{"value":110,"modifier":"EXACT"},"end":{"value":117,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]},{"type":"Disulfide bond","location":{"start":{"value":280,"modifier":"EXACT"},"end":{"value":284,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR633144-2"}]},{"type":"Disulfide bond","location":{"start":{"value":323,"modifier":"EXACT"},"end":{"value":360,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"},{"evidenceCode":"ECO:0000256","source":"PIRSR","id":"PIRSR601461-2"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0A1B0GV23"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0A1B0GV23"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490344.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490344.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490344.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"21269460","citationType":"journal article","authors":["Burkard T.R.","Planyavsky M.","Kaupe I.","Breitwieser F.P.","Burckstummer T.","Bennett K.L.","Superti-Furga G.","Colinge J."],"citationCrossReferences":[{"database":"PubMed","id":"21269460"},{"database":"DOI","id":"10.1186/1752-0509-5-17"}],"title":"Initial characterization of the human central proteome.","publicationDate":"2011","journal":"BMC Syst. Biol.","firstPage":"17","lastPage":"17","volume":"5"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"21269460"}]},{"referenceNumber":5,"citation":{"id":"25944712","citationType":"journal article","authors":["Vaca Jacome A.S.","Rabilloud T.","Schaeffer-Reiss C.","Rompais M.","Ayoub D.","Lane L.","Bairoch A.","Van Dorsselaer A.","Carapito C."],"citationCrossReferences":[{"database":"PubMed","id":"25944712"}],"title":"N-terminome analysis of the human mitochondrial proteome.","publicationDate":"2015","journal":"Proteomics","firstPage":"2519","lastPage":"2524","volume":"15"},"referencePositions":["IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]"],"evidences":[{"evidenceCode":"ECO:0007829","source":"PubMed","id":"25944712"}]},{"referenceNumber":6,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000490344.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"AlphaFoldDB","id":"A0A1B0GV23","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"A0A1B0GV23","properties":[{"key":"Description","value":"-"}]},{"database":"GlyCosmos","id":"A0A1B0GV23","properties":[{"key":"glycosylation","value":"2 sites, No reported glycans"}]},{"database":"MassIVE","id":"A0A1B0GV23","properties":[{"key":"Description","value":"-"}]},{"database":"Antibodypedia","id":"880","properties":[{"key":"antibodies","value":"1212 antibodies from 49 providers"}]},{"database":"Ensembl","id":"ENST00000637815.2","properties":[{"key":"ProteinId","value":"ENSP00000490344.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"VEuPathDB","id":"HostDB:ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"ChiTaRS","id":"CTSD","properties":[{"key":"OrganismName","value":"human"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"Bgee","id":"ENSG00000117984","properties":[{"key":"ExpressionPatterns","value":"Expressed in right adrenal gland cortex and 199 other cell types or tissues"}]},{"database":"ExpressionAtlas","id":"A0A1B0GV23","properties":[{"key":"ExpressionPatterns","value":"baseline and differential"}]},{"database":"GO","id":"GO:0005764","properties":[{"key":"GoTerm","value":"C:lysosome"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-SubCell"}]},{"database":"GO","id":"GO:0004190","properties":[{"key":"GoTerm","value":"F:aspartic-type endopeptidase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-EC"}]},{"database":"GO","id":"GO:0006508","properties":[{"key":"GoTerm","value":"P:proteolysis"},{"key":"GoEvidenceType","value":"IEA:InterPro"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":406,"molWeight":43831,"crc64":"3EF793C92F82F143","md5":"9E84185BAAEDD291AB3B6A78413DCD97"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2,"Glycosylation":2,"Disulfide bond":4},"uniParcId":"UPI0007E52C21"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0ACI8VAQ0","uniProtkbId":"A0ACI8VAQ0_HUMAN","entryAudit":{"firstPublicDate":"2026-06-10","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2026-06-10","entryVersion":2,"sequenceVersion":1},"annotationScore":2.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632505.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632505.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":467,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5051204908"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":462,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Region","location":{"start":{"value":267,"modifier":"EXACT"},"end":{"value":296,"modifier":"EXACT"}},"description":"Disordered","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"MobiDB-lite"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":350,"modifier":"EXACT"},"end":{"value":350,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0ACI8VAQ0"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0ACI8VAQ0"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022729"},{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632505.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632505.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632505.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632505.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"Ensembl","id":"ENST00000962446.1","properties":[{"key":"ProteinId","value":"ENSP00000632505.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"OpenTargets","id":"ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000066","properties":[{"key":"EntryName","value":"Napsin A aspartic peptidase"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"2"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRIPAVGLAGPLCSAPCLGSEYSLATPRAKLLTRGQDTGEPAGPFPQEPWGTGEAARDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":467,"molWeight":50098,"crc64":"281F9496692EB4B1","md5":"4984AFB5B497312A3128B44C31BECC4D"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Region":1,"Active site":2},"uniParcId":"UPI0040321A90"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0ACI8RMI1","uniProtkbId":"A0ACI8RMI1_HUMAN","entryAudit":{"firstPublicDate":"2026-06-10","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2026-06-10","entryVersion":2,"sequenceVersion":1},"annotationScore":2.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577883.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577883.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":395,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5051294661"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":390,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":295,"modifier":"EXACT"},"end":{"value":295,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0ACI8RMI1"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0ACI8RMI1"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577883.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577883.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577883.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577883.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"Ensembl","id":"ENST00000907824.1","properties":[{"key":"ProteinId","value":"ENSP00000577883.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"OpenTargets","id":"ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"3"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":395,"molWeight":42749,"crc64":"54895DD3DED65E66","md5":"6E36D5AA52BCE94144261C77CAF627CF"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2},"uniParcId":"UPI0040317751"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0ACI8STF4","uniProtkbId":"A0ACI8STF4_HUMAN","entryAudit":{"firstPublicDate":"2026-06-10","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2026-06-10","entryVersion":2,"sequenceVersion":1},"annotationScore":2.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000586429.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000586429.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":410,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5051420777"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":405,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":293,"modifier":"EXACT"},"end":{"value":293,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0ACI8STF4"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0ACI8STF4"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000586429.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000586429.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000586429.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000586429.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"Ensembl","id":"ENST00000916370.1","properties":[{"key":"ProteinId","value":"ENSP00000586429.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"OpenTargets","id":"ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":410,"molWeight":44366,"crc64":"1FC36D5828013E87","md5":"61FED45B413179B3E5CCA45C0436D8F4"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2},"uniParcId":"UPI00403159EF"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0ACI8VAS8","uniProtkbId":"A0ACI8VAS8_HUMAN","entryAudit":{"firstPublicDate":"2026-06-10","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2026-06-10","entryVersion":2,"sequenceVersion":1},"annotationScore":2.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632504.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632504.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":406,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5051357869"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":401,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":295,"modifier":"EXACT"},"end":{"value":295,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0ACI8VAS8"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0ACI8VAS8"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632504.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632504.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632504.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632504.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"Ensembl","id":"ENST00000962445.1","properties":[{"key":"ProteinId","value":"ENSP00000632504.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"OpenTargets","id":"ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":406,"molWeight":43815,"crc64":"B99513602931C03A","md5":"A076836304D33AE332FE22A60F6F683E"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2},"uniParcId":"UPI0040319FA6"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0ACI8R5F3","uniProtkbId":"A0ACI8R5F3_HUMAN","entryAudit":{"firstPublicDate":"2026-06-10","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2026-06-10","entryVersion":2,"sequenceVersion":1},"annotationScore":2.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577884.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577884.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":371,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5051378102"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":366,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":254,"modifier":"EXACT"},"end":{"value":254,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0ACI8R5F3"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0ACI8R5F3"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022729"},{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577884.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577884.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577884.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577884.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"Ensembl","id":"ENST00000907825.1","properties":[{"key":"ProteinId","value":"ENSP00000577884.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"OpenTargets","id":"ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000066","properties":[{"key":"EntryName","value":"Napsin A aspartic peptidase"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.60.40.1960","properties":[{"key":"EntryName","value":"-"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":371,"molWeight":39967,"crc64":"C721D3DBF0F285D7","md5":"146FFCDBBA06C0955A9A80D2C5554FD7"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2},"uniParcId":"UPI0040322E2B"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0ACI8V6H6","uniProtkbId":"A0ACI8V6H6_HUMAN","entryAudit":{"firstPublicDate":"2026-06-10","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2026-06-10","entryVersion":2,"sequenceVersion":1},"annotationScore":2.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632506.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632506.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":427,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5051441727"},{"type":"Domain","location":{"start":{"value":94,"modifier":"EXACT"},"end":{"value":422,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":112,"modifier":"EXACT"},"end":{"value":112,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":310,"modifier":"EXACT"},"end":{"value":310,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0ACI8V6H6"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0ACI8V6H6"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632506.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632506.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632506.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632506.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"Ensembl","id":"ENST00000962447.1","properties":[{"key":"ProteinId","value":"ENSP00000632506.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"OpenTargets","id":"ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDPLLCPLLLQGEAGSKAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":427,"molWeight":46073,"crc64":"5DCA2F95E64D6B6A","md5":"5D731C1E56FA0340F3DDCC6590EBCB91"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2},"uniParcId":"UPI004031A7F2"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0ACI8RJ76","uniProtkbId":"A0ACI8RJ76_HUMAN","entryAudit":{"firstPublicDate":"2026-06-10","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2026-06-10","entryVersion":2,"sequenceVersion":1},"annotationScore":2.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577882.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577882.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":168,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5051243805"},{"type":"Domain","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":163,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":51,"modifier":"EXACT"},"end":{"value":51,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0ACI8RJ76"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0ACI8RJ76"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577882.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577882.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577882.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577882.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"Ensembl","id":"ENST00000907823.1","properties":[{"key":"ProteinId","value":"ENSP00000577882.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"OpenTargets","id":"ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"FunFam","id":"2.40.70.10:FF:000044","properties":[{"key":"EntryName","value":"Lysosomal aspartic protease"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":168,"molWeight":18036,"crc64":"2033284E992C65DD","md5":"DADF0C6CD454759816D00ABAB0CB86B8"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":1},"uniParcId":"UPI0040325808"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0ACI8SE50","uniProtkbId":"A0ACI8SE50_HUMAN","entryAudit":{"firstPublicDate":"2026-06-10","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2026-06-10","entryVersion":2,"sequenceVersion":1},"annotationScore":2.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577885.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577885.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":313,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5051463812"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":313,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":196,"modifier":"EXACT"},"end":{"value":196,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0ACI8SE50"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0ACI8SE50"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577885.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577885.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577885.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000577885.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"Ensembl","id":"ENST00000907826.1","properties":[{"key":"ProteinId","value":"ENSP00000577885.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"OpenTargets","id":"ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000002","properties":[{"key":"EntryName","value":"Vacuolar aspartic proteinase"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSVPCQSASSASALGGVKVERQVFGEATKQPGITFIAAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":313,"molWeight":33502,"crc64":"74879FA22A6F4A85","md5":"5110DBFEC79C4CC2109600E7A3D74BB2"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2},"uniParcId":"UPI00403223BE"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0ACI8VBV2","uniProtkbId":"A0ACI8VBV2_HUMAN","entryAudit":{"firstPublicDate":"2026-06-10","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2026-06-10","entryVersion":2,"sequenceVersion":1},"annotationScore":2.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632503.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"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:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"Pepsin A"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}],"value":"3.4.23.1"}]}},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632503.1"}],"value":"CTSD"}}],"comments":[{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Preferential cleavage: hydrophobic, preferably aromatic, residues in P1 and P1' positions. Cleaves 1-Phe-|-Val-2, 4-Gln-|-His-5, 13-Glu-|-Ala-14, 14-Ala-|-Leu-15, 15-Leu-|-Tyr-16, 16-Tyr-|-Leu-17, 23-Gly-|-Phe-24, 24-Phe-|-Phe-25 and 25-Phe-|-Tyr-26 bonds in the B chain of insulin.","ecNumber":"3.4.23.1","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"Specificity similar to, but narrower than, that of pepsin A. Does not cleave the 4-Gln-|-His-5 bond in B chain of insulin.","ecNumber":"3.4.23.5","evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00000585"}]}},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00004371"}],"value":"Lysosome","id":"SL-0158"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"},{"evidenceCode":"ECO:0000256","source":"RuleBase","id":"RU000454"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":408,"modifier":"EXACT"}},"description":"Pepsin A","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5051403930"},{"type":"Domain","location":{"start":{"value":79,"modifier":"EXACT"},"end":{"value":403,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Active site","location":{"start":{"value":97,"modifier":"EXACT"},"end":{"value":97,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]},{"type":"Active site","location":{"start":{"value":291,"modifier":"EXACT"},"end":{"value":291,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU10094"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023180"}],"id":"KW-0325","category":"PTM","name":"Glycoprotein"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023228"}],"id":"KW-0458","category":"Cellular component","name":"Lysosome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0007829","source":"PeptideAtlas","id":"A0ACI8VBV2"},{"evidenceCode":"ECO:0007829","source":"ProteomicsDB","id":"A0ACI8VBV2"}],"id":"KW-1267","category":"Technical term","name":"Proteomics identification"},{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632503.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632503.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632503.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000632503.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"Ensembl","id":"ENST00000962444.1","properties":[{"key":"ProteinId","value":"ENSP00000632503.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"OpenTargets","id":"ENSG00000117984","properties":[{"key":"Description","value":"-"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"CDD","id":"cd05490","properties":[{"key":"EntryName","value":"Cathepsin_D2"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000039","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"2.40.70.10:FF:000047","properties":[{"key":"EntryName","value":"Cathepsin D preproprotein"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"2"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR001969","properties":[{"key":"EntryName","value":"Aspartic_peptidase_AS"}]},{"database":"InterPro","id":"IPR012848","properties":[{"key":"EntryName","value":"Aspartic_peptidase_N"}]},{"database":"InterPro","id":"IPR033144","properties":[{"key":"EntryName","value":"Cathepsin_D"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF07966","properties":[{"key":"EntryName","value":"A1_Propeptide"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"PRINTS","id":"PR00792","properties":[{"key":"EntryName","value":"PEPSIN"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS00141","properties":[{"key":"EntryName","value":"ASP_PROTEASE"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVRIPLHKFTSIRRTMSEVGGSVEDLIAKGPVSKYSQAVPAVTEGPIPEVLKNYMDAQYYGEIGIGTPPQCFTVVFDTGSSNLWVPSIHCKLLDIACWIHHKYNSDKSSTYVKNGTSFDIHYGSGSLSGYLSQDTVSSASSASALGGVKVERQVFGEATKQPGITFIAAKFDGILGMAYPRISVNNVLPVFDNLMQQKLVDQNIFSFYLSRDPDAQPGGELMLGGTDSKYYKGSLSYLNVTRKAYWQVHLDQVEVASGLTLCKEGCEAIVDTGTSLMVGPVDEVRELQKAIGAVPLIQGEYMIPCEKVSTLPAITLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":408,"molWeight":44125,"crc64":"DCAA7455693719DA","md5":"233DE9D844368D74722110FB2B0AE5A9"},"extraAttributes":{"countByCommentType":{"CATALYTIC ACTIVITY":2,"SUBCELLULAR LOCATION":1,"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Signal":1,"Chain":1,"Domain":1,"Active site":2},"uniParcId":"UPI0040322901"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0A7I2V2N3","uniProtkbId":"A0A7I2V2N3_HUMAN","entryAudit":{"firstPublicDate":"2021-04-07","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2021-04-07","entryVersion":23,"sequenceVersion":1},"annotationScore":1.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000503019.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"lineage":["Eukaryota","Metazoa","Chordata","Craniata","Vertebrata","Euteleostomi","Mammalia","Eutheria","Euarchontoglires","Primates","Haplorrhini","Catarrhini","Hominidae","Homo"]},"proteinExistence":"4: Predicted","proteinDescription":{"submissionNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000503019.1"}],"value":"Cathepsin D"}}]},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000503019.1"}],"value":"CTSD"}}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":20,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}]},{"type":"Chain","location":{"start":{"value":21,"modifier":"EXACT"},"end":{"value":93,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"featureId":"PRO_5029746579"}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}],"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"SAM","id":"SignalP"}],"id":"KW-0732","category":"Domain","name":"Signal"}],"references":[{"referenceNumber":1,"citation":{"id":"11237011","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"authors":["Lander E.S.","Linton L.M.","Birren B.","Nusbaum C.","Zody M.C.","Baldwin J.","Devon K.","Dewar K.","Doyle M.","FitzHugh W.","Funke R.","Gage D.","Harris K.","Heaford A.","Howland J.","Kann L.","Lehoczky J.","LeVine R.","McEwan P.","McKernan K.","Meldrim J.","Mesirov J.P.","Miranda C.","Morris W.","Naylor J.","Raymond C.","Rosetti M.","Santos R.","Sheridan A.","Sougnez C.","Stange-Thomann N.","Stojanovic N.","Subramanian A.","Wyman D.","Rogers J.","Sulston J.","Ainscough R.","Beck S.","Bentley D.","Burton J.","Clee C.","Carter N.","Coulson A.","Deadman R.","Deloukas P.","Dunham A.","Dunham I.","Durbin R.","French L.","Grafham D.","Gregory S.","Hubbard T.","Humphray S.","Hunt A.","Jones M.","Lloyd C.","McMurray A.","Matthews L.","Mercer S.","Milne S.","Mullikin J.C.","Mungall A.","Plumb R.","Ross M.","Shownkeen R.","Sims S.","Waterston R.H.","Wilson R.K.","Hillier L.W.","McPherson J.D.","Marra M.A.","Mardis E.R.","Fulton L.A.","Chinwalla A.T.","Pepin K.H.","Gish W.R.","Chissoe S.L.","Wendl M.C.","Delehaunty K.D.","Miner T.L.","Delehaunty A.","Kramer J.B.","Cook L.L.","Fulton R.S.","Johnson D.L.","Minx P.J.","Clifton S.W.","Hawkins T.","Branscomb E.","Predki P.","Richardson P.","Wenning S.","Slezak T.","Doggett N.","Cheng J.F.","Olsen A.","Lucas S.","Elkin C.","Uberbacher E.","Frazier M.","Gibbs R.A.","Muzny D.M.","Scherer S.E.","Bouck J.B.","Sodergren E.J.","Worley K.C.","Rives C.M.","Gorrell J.H.","Metzker M.L.","Naylor S.L.","Kucherlapati R.S.","Nelson D.L.","Weinstock G.M.","Sakaki Y.","Fujiyama A.","Hattori M.","Yada T.","Toyoda A.","Itoh T.","Kawagoe C.","Watanabe H.","Totoki Y.","Taylor T.","Weissenbach J.","Heilig R.","Saurin W.","Artiguenave F.","Brottier P.","Bruls T.","Pelletier E.","Robert C.","Wincker P.","Smith D.R.","Doucette-Stamm L.","Rubenfield M.","Weinstock K.","Lee H.M.","Dubois J.","Rosenthal A.","Platzer M.","Nyakatura G.","Taudien S.","Rump A.","Yang H.","Yu J.","Wang J.","Huang G.","Gu J.","Hood L.","Rowen L.","Madan A.","Qin S.","Davis R.W.","Federspiel N.A.","Abola A.P.","Proctor M.J.","Myers R.M.","Schmutz J.","Dickson M.","Grimwood J.","Cox D.R.","Olson M.V.","Kaul R.","Raymond C.","Shimizu N.","Kawasaki K.","Minoshima S.","Evans G.A.","Athanasiou M.","Schultz R.","Roe B.A.","Chen F.","Pan H.","Ramser J.","Lehrach H.","Reinhardt R.","McCombie W.R.","de la Bastide M.","Dedhia N.","Blocker H.","Hornischer K.","Nordsiek G.","Agarwala R.","Aravind L.","Bailey J.A.","Bateman A.","Batzoglou S.","Birney E.","Bork P.","Brown D.G.","Burge C.B.","Cerutti L.","Chen H.C.","Church D.","Clamp M.","Copley R.R.","Doerks T.","Eddy S.R.","Eichler E.E.","Furey T.S.","Galagan J.","Gilbert J.G.","Harmon C.","Hayashizaki Y.","Haussler D.","Hermjakob H.","Hokamp K.","Jang W.","Johnson L.S.","Jones T.A.","Kasif S.","Kaspryzk A.","Kennedy S.","Kent W.J.","Kitts P.","Koonin E.V.","Korf I.","Kulp D.","Lancet D.","Lowe T.M.","McLysaght A.","Mikkelsen T.","Moran J.V.","Mulder N.","Pollara V.J.","Ponting C.P.","Schuler G.","Schultz J.","Slater G.","Smit A.F.","Stupka E.","Szustakowski J.","Thierry-Mieg D.","Thierry-Mieg J.","Wagner L.","Wallis J.","Wheeler R.","Williams A.","Wolf Y.I.","Wolfe K.H.","Yang S.P.","Yeh R.F.","Collins F.","Guyer M.S.","Peterson J.","Felsenfeld A.","Wetterstrand K.A.","Patrinos A.","Morgan M.J.","de Jong P.","Catanese J.J.","Osoegawa K.","Shizuya H.","Choi S.","Chen Y.J."],"citationCrossReferences":[{"database":"PubMed","id":"11237011"},{"database":"DOI","id":"10.1038/35057062"}],"title":"Initial sequencing and analysis of the human genome.","publicationDate":"2001","journal":"Nature","firstPage":"860","lastPage":"921","volume":"409"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000503019.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":2,"citation":{"id":"15496913","citationType":"journal article","authoringGroup":["International Human Genome Sequencing Consortium"],"citationCrossReferences":[{"database":"PubMed","id":"15496913"},{"database":"DOI","id":"10.1038/nature03001"}],"title":"Finishing the euchromatic sequence of the human genome.","publicationDate":"2004","journal":"Nature","firstPage":"931","lastPage":"945","volume":"431"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000503019.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":3,"citation":{"id":"16554811","citationType":"journal article","authors":["Taylor T.D.","Noguchi H.","Totoki Y.","Toyoda A.","Kuroki Y.","Dewar K.","Lloyd C.","Itoh T.","Takeda T.","Kim D.W.","She X.","Barlow K.F.","Bloom T.","Bruford E.","Chang J.L.","Cuomo C.A.","Eichler E.","FitzGerald M.G.","Jaffe D.B.","LaButti K.","Nicol R.","Park H.S.","Seaman C.","Sougnez C.","Yang X.","Zimmer A.R.","Zody M.C.","Birren B.W.","Nusbaum C.","Fujiyama A.","Hattori M.","Rogers J.","Lander E.S.","Sakaki Y."],"citationCrossReferences":[{"database":"PubMed","id":"16554811"},{"database":"DOI","id":"10.1038/nature04632"}],"title":"Human chromosome 11 DNA sequence and analysis including novel gene identification.","publicationDate":"2006","journal":"Nature","firstPage":"497","lastPage":"500","volume":"440"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000503019.1"},{"evidenceCode":"ECO:0000313","source":"Proteomes","id":"UP000005640"}]},{"referenceNumber":4,"citation":{"id":"CI-20IM145UGQVF7","citationType":"submission","authoringGroup":["Ensembl"],"publicationDate":"JAN-2026","submissionDatabase":"UniProtKB"},"referencePositions":["IDENTIFICATION"],"evidences":[{"evidenceCode":"ECO:0000313","source":"Ensembl","id":"ENSP00000503019.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"AC068580","properties":[{"key":"ProteinId","value":"-"},{"key":"Status","value":"NOT_ANNOTATED_CDS"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"AlphaFoldDB","id":"A0A7I2V2N3","properties":[{"key":"Description","value":"-"}]},{"database":"Ensembl","id":"ENST00000678991.1","properties":[{"key":"ProteinId","value":"ENSP00000503019.1"},{"key":"GeneId","value":"ENSG00000117984.16"}]},{"database":"HGNC","id":"HGNC:2529","properties":[{"key":"GeneName","value":"CTSD"}]},{"database":"GeneTree","id":"ENSGT00940000155733","properties":[{"key":"Description","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000005640","properties":[{"key":"Component","value":"Chromosome 11"}]},{"database":"Bgee","id":"ENSG00000117984","properties":[{"key":"ExpressionPatterns","value":"Expressed in right adrenal gland cortex and 199 other cell types or tissues"}]},{"database":"ExpressionAtlas","id":"A0A7I2V2N3","properties":[{"key":"ExpressionPatterns","value":"baseline and differential"}]}],"sequence":{"value":"MQPSSLLPLALCLLAAPASALVSFQVTGRSLVLAVATLRFPRGLQVDVQHWQRRPAGGRASLICPDPWLTGSRCTSSRPSAGPCRRLGALWRT","length":93,"molWeight":9978,"crc64":"1F11CD8C680C9B11","md5":"7D37748070FFCBF8E2D24CD438A63DBC"},"extraAttributes":{"countByFeatureType":{"Signal":1,"Chain":1},"uniParcId":"UPI0018BDCB7F"}},{"entryType":"UniProtKB unreviewed (TrEMBL)","primaryAccession":"A0A0S2Z3L4","uniProtkbId":"A0A0S2Z3L4_HUMAN","entryAudit":{"firstPublicDate":"2016-02-17","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2016-02-17","entryVersion":34,"sequenceVersion":1},"annotationScore":1.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"ALQ33466.1"}],"lineage":["Eukaryota","Metazoa","Chordata","Craniata","Vertebrata","Euteleostomi","Mammalia","Eutheria","Euarchontoglires","Primates","Haplorrhini","Catarrhini","Hominidae","Homo"]},"proteinExistence":"2: Evidence at transcript level","proteinDescription":{"submissionNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"ALQ33466.1"}],"value":"Cathepsin D isoform 2"}}],"flag":"Fragment"},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"ALQ33466.1"}],"value":"CTSD"}}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00007447"},{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Belongs to the peptidase A1 family"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"PROSITE-ProRule","id":"PRU01103"}],"value":"Lacks conserved residue(s) required for the propagation of feature annotation"}],"commentType":"CAUTION"}],"features":[{"type":"Domain","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":78,"modifier":"EXACT"}},"description":"Peptidase A1","evidences":[{"evidenceCode":"ECO:0000259","source":"PROSITE","id":"PS51767"}]},{"type":"Non-terminal residue","location":{"start":{"value":83,"modifier":"EXACT"},"end":{"value":83,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"ALQ33466.1"}]}],"keywords":[{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022750"}],"id":"KW-0064","category":"Molecular function","name":"Aspartyl protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023157"}],"id":"KW-1015","category":"PTM","name":"Disulfide bond"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022801"}],"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00022670"}],"id":"KW-0645","category":"Molecular function","name":"Protease"},{"evidences":[{"evidenceCode":"ECO:0000256","source":"ARBA","id":"ARBA00023145"}],"id":"KW-0865","category":"PTM","name":"Zymogen"}],"references":[{"referenceNumber":1,"citation":{"id":"26871637","citationType":"journal article","authors":["Yang X.","Coulombe-Huntington J.","Kang S.","Sheynkman G.M.","Hao T.","Richardson A.","Sun S.","Yang F.","Shen Y.A.","Murray R.","Spirohn K.","Begg B.E.","Duran-Frigola M.","MacWilliams A.","Pevzner S.J.","Zhong Q.","Trigg S.A.","Tam S.","Ghamsari L.","Sahni N.","Yi S.","Rodriguez M.D.","Balcha D.","Tan G.","Costanzo M.","Andrews B.","Boone C.","Zhou X.J.","Salehi-Ashtiani K.","Charloteaux B.","Chen A.","Calderwood M.A.","Aloy P.","Roth F.P.","Hill D.E.","Iakoucheva L.M.","Xia Y.","Vidal M."],"citationCrossReferences":[{"database":"PubMed","id":"26871637"},{"database":"DOI","id":"10.1016/j.cell.2016.01.029"}],"title":"Widespread Expansion of Protein Interaction Capabilities by Alternative Splicing.","publicationDate":"2016","journal":"Cell","firstPage":"805","lastPage":"817","volume":"164"},"referencePositions":["NUCLEOTIDE SEQUENCE"],"evidences":[{"evidenceCode":"ECO:0000313","source":"EMBL","id":"ALQ33466.1"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"KU178008","properties":[{"key":"ProteinId","value":"ALQ33466.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"mRNA"}]},{"database":"AlphaFoldDB","id":"A0A0S2Z3L4","properties":[{"key":"Description","value":"-"}]},{"database":"PeptideAtlas","id":"A0A0S2Z3L4","properties":[{"key":"Description","value":"-"}]},{"database":"OrthoDB","id":"771136at2759","properties":[{"key":"Description","value":"-"}]},{"database":"ChiTaRS","id":"CTSD","properties":[{"key":"OrganismName","value":"human"}]},{"database":"GO","id":"GO:0004190","properties":[{"key":"GoTerm","value":"F:aspartic-type endopeptidase activity"},{"key":"GoEvidenceType","value":"IEA:InterPro"}]},{"database":"GO","id":"GO:0006508","properties":[{"key":"GoTerm","value":"P:proteolysis"},{"key":"GoEvidenceType","value":"IEA:InterPro"}]},{"database":"FunFam","id":"2.40.70.10:FF:000044","properties":[{"key":"EntryName","value":"Lysosomal aspartic protease"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"2.40.70.10","properties":[{"key":"EntryName","value":"Acid Proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"InterPro","id":"IPR001461","properties":[{"key":"EntryName","value":"Aspartic_peptidase_A1"}]},{"database":"InterPro","id":"IPR033121","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"}]},{"database":"InterPro","id":"IPR021109","properties":[{"key":"EntryName","value":"Peptidase_aspartic_dom_sf"}]},{"database":"PANTHER","id":"PTHR47966","properties":[{"key":"EntryName","value":"BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47966:SF42","properties":[{"key":"EntryName","value":"CATHEPSIN D"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF00026","properties":[{"key":"EntryName","value":"Asp"},{"key":"MatchStatus","value":"1"}]},{"database":"SUPFAM","id":"SSF50630","properties":[{"key":"EntryName","value":"Acid proteases"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS51767","properties":[{"key":"EntryName","value":"PEPTIDASE_A1"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MQPSSLLPLTLKLGGKGYKLSPEDYTLKVSQAGKTLCLSGFMGMDIPPPSGPLWILGDVFIGRYYTVFDRDNNRVGFAEAARL","length":83,"molWeight":9082,"crc64":"089CF45B8A74F097","md5":"24194869A93167CD39C5D185E0762411"},"extraAttributes":{"countByCommentType":{"SIMILARITY":1,"CAUTION":1},"countByFeatureType":{"Domain":1,"Non-terminal residue":1},"uniParcId":"UPI00071F5381"}}]}