{"results":[{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"P03950","secondaryAccessions":["Q05CV1","Q53X86","Q6P5T2","Q8WXE7"],"uniProtkbId":"ANGI_HUMAN","entryAudit":{"firstPublicDate":"1986-10-23","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"1986-10-23","entryVersion":255,"sequenceVersion":1},"annotationScore":5.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"lineage":["Eukaryota","Metazoa","Chordata","Craniata","Vertebrata","Euteleostomi","Mammalia","Eutheria","Euarchontoglires","Primates","Haplorrhini","Catarrhini","Hominidae","Homo"]},"proteinExistence":"1: Evidence at protein level","proteinDescription":{"recommendedName":{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"2866795"}],"value":"Angiogenin"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"1400510"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21855800"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2424496"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2459697"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"28176817"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38718836"}],"value":"3.1.27.-"}]},"alternativeNames":[{"fullName":{"value":"Ribonuclease 5"},"shortNames":[{"value":"RNase 5"}]}],"flag":"Precursor"},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"11919285"},{"evidenceCode":"ECO:0000312","source":"HGNC","id":"HGNC:483"}],"value":"ANG"},"synonyms":[{"value":"RNASE5"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P21570"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12051708"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"1400510"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15735021"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19332886"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19354288"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20129916"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21855800"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23047679"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23843625"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2424496"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2459697"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2730651"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27518564"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"28176817"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29100074"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29748193"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"3122207"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"31582561"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"32510170"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"3289612"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38718836"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"4074709"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8127865"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8159680"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8448182"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8570639"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8622921"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9578571"}],"value":"Secreted ribonuclease that can either promote or restrict cell proliferation of target cells, depending on the context (PubMed:12051708, PubMed:1400510, PubMed:19332886, PubMed:20129916, PubMed:21855800, PubMed:23047679, PubMed:23843625, PubMed:2424496, PubMed:2459697, PubMed:2730651, PubMed:27518564, PubMed:28176817, PubMed:29100074, PubMed:29748193, PubMed:3122207, PubMed:32510170, PubMed:38718836, PubMed:8159680, PubMed:8570639, PubMed:8622921, PubMed:9578571). Endocytosed in target cells via its receptor PLXNB2 and translocates to the cytoplasm or nucleus (PubMed:29100074, PubMed:32510170). Under stress conditions, localizes to the cytoplasm and promotes the assembly of stress granules (SGs): specifically cleaves a subset of tRNAs within anticodon loops to produce tRNA-derived stress-induced fragments (tiRNAs), resulting in translation repression and inhibition of cell proliferation (PubMed:1400510, PubMed:19332886, PubMed:20129916, PubMed:21855800, PubMed:23047679, PubMed:27518564, PubMed:29100074, PubMed:29748193, PubMed:32510170, PubMed:38718836). tiRNas also prevent formation of apoptosome, thereby promoting cell survival (By similarity). Preferentially cleaves RNAs between a pyrimidine and an adenosine residue, suggesting that it cleaves the anticodon loop of tRNA(Ala) (32-UUAGCAU-38) after positions 33 and 36 (PubMed:3289612, PubMed:38718836). Cleaves a subset of tRNAs, including tRNA(Ala), tRNA(Glu), tRNA(Gly), tRNA(Lys), tRNA(Val), tRNA(His), tRNA(Asp) and tRNA(Sec) (PubMed:31582561). Under growth conditions and in differentiated cells, translocates to the nucleus and stimulates ribosomal RNA (rRNA) transcription, including that containing the initiation site sequences of 45S rRNA, thereby promoting cell growth and proliferation (PubMed:12051708, PubMed:15735021, PubMed:27518564, PubMed:29100074, PubMed:8127865). Angiogenin induces vascularization of normal and malignant tissues via its ability to promote rRNA transcription (PubMed:19354288, PubMed:4074709, PubMed:8448182). Involved in hematopoietic stem and progenitor cell (HSPC) growth and survival by promoting rRNA transcription in growth conditions and inhibiting translation in response to stress, respectively (PubMed:27518564). Mediates the crosstalk between myeloid and intestinal epithelial cells to protect the intestinal epithelial barrier integrity: secreted by myeloid cells and promotes intestinal epithelial cells proliferation and survival (PubMed:32510170). Also mediates osteoclast-endothelial cell crosstalk in growing bone: produced by osteoclasts and protects the neighboring vascular cells against senescence by promoting rRNA transcription (By similarity)"}],"commentType":"FUNCTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19332886"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19354288"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23843625"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"3243277"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"32510170"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38718836"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8159680"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8622921"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9578571"}],"value":"Has weak tRNA ribonuclease activity by itself due to partial autoinhibition by its C-terminus (residues 140-147), which folds into a short alpha-helix that partially occludes the substrate-binding site (PubMed:38718836, PubMed:8159680, PubMed:8622921, PubMed:9578571). In absence of stress, the ribonuclease activity is inhibited by RNH1 in the cytoplasm (PubMed:19332886, PubMed:19354288, PubMed:23843625, PubMed:3243277, PubMed:32510170, PubMed:38718836). In response to stress, dissociates from RNH1 in the cytoplasm and associates with cytoplasmic ribosomes with vacant A-sites: ribosomes directly activate the tRNA ribonuclease activity of ANG by refolding the C-terminal alpha-helix (PubMed:38718836). In response to stress, the angiogenic activity of ANG is inhibited by RNH1 in the nucleus (PubMed:23843625)"}],"commentType":"ACTIVITY REGULATION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10413501"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19332886"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19354288"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23843625"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25372031"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27518564"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"3243277"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"32510170"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"37218877"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9311977"}],"value":"Homodimer (PubMed:25372031). Interacts with RNH1; inhibiting ANG ribonuclease activity (PubMed:10413501, PubMed:19332886, PubMed:19354288, PubMed:23843625, PubMed:27518564, PubMed:3243277, PubMed:32510170, PubMed:9311977). Interacts with PCNA (PubMed:37218877)"}],"commentType":"SUBUNIT"},{"commentType":"INTERACTION","interactions":[{"interactantOne":{"uniProtKBAccession":"P03950","intActId":"EBI-525291"},"interactantTwo":{"uniProtKBAccession":"P35609","geneName":"ACTN2","intActId":"EBI-77797"},"numberOfExperiments":4,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P03950","intActId":"EBI-525291"},"interactantTwo":{"uniProtKBAccession":"P19883","geneName":"FST","intActId":"EBI-1571188"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P03950","intActId":"EBI-525291"},"interactantTwo":{"uniProtKBAccession":"P12004","geneName":"PCNA","intActId":"EBI-358311"},"numberOfExperiments":4,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P03950","intActId":"EBI-525291"},"interactantTwo":{"uniProtKBAccession":"Q03405","geneName":"PLAUR","intActId":"EBI-716505"},"numberOfExperiments":5,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P03950","intActId":"EBI-525291"},"interactantTwo":{"uniProtKBAccession":"P13489","geneName":"RNH1","intActId":"EBI-1237106"},"numberOfExperiments":6,"organismDiffer":false}]},{"commentType":"SUBCELLULAR LOCATION","note":{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12051708"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29100074"}],"value":"The secreted protein is rapidly endocytosed by target cells following interaction with PLXNB2 receptor and translocated to the cytoplasm and nucleus (PubMed:29100074). In the nucleus, accumulates in the nucleolus and binds to DNA (PubMed:12051708)"}]},"subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23047679"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29748193"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"3663649"}],"value":"Secreted","id":"SL-0243"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12051708"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15735021"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23047679"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23843625"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25372031"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29100074"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8127865"}],"value":"Nucleus","id":"SL-0191"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12051708"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"7945327"}],"value":"Nucleus, nucleolus","id":"SL-0188"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23843625"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27518564"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29100074"}],"value":"Cytoplasm, Stress granule","id":"SL-0496"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17886298"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2440105"}],"value":"Expressed predominantly in the liver (PubMed:2440105). Also detected in endothelial cells and spinal cord neurons (PubMed:17886298, PubMed:2440105)"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2440105"}],"value":"Low level expression in the developing fetus, increased in the neonate, and maximal in the adult"}],"commentType":"DEVELOPMENTAL STAGE"},{"commentType":"DISEASE","disease":{"diseaseId":"Amyotrophic lateral sclerosis 9","diseaseAccession":"DI-00113","acronym":"ALS9","description":"A neurodegenerative disorder affecting upper motor neurons in the brain and lower motor neurons in the brain stem and spinal cord, resulting in fatal paralysis. Sensory abnormalities are absent. The pathologic hallmarks of the disease include pallor of the corticospinal tract due to loss of motor neurons, presence of ubiquitin-positive inclusions within surviving motor neurons, and deposition of pathologic aggregates. The etiology of amyotrophic lateral sclerosis is likely to be multifactorial, involving both genetic and environmental factors. The disease is inherited in 5-10% of the cases.","diseaseCrossReference":{"database":"MIM","id":"611895"},"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15557516"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16501576"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17703939"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17886298"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17900154"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18087731"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18852347"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19153377"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19332886"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19363631"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22190368"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22292798"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22292843"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25372031"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29748193"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38718836"}]},"note":{"texts":[{"value":"Disease susceptibility is associated with variants affecting the gene represented in this entry"}]}},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Belongs to the pancreatic ribonuclease family"}],"commentType":"SIMILARITY"},{"commentType":"SEQUENCE CAUTION","sequenceCautionType":"Erroneous initiation","sequence":"AAH20704.1","note":"Truncated N-terminus.","evidences":[{"evidenceCode":"ECO:0000305"}]},{"commentType":"WEB RESOURCE","resourceName":"Functional Glycomics Gateway - Glycan Binding","resourceUrl":"http://www.functionalglycomics.org/glycomics/GBPServlet?&operationType=view&cbpId=cbp_hum_other_803","ftp":false,"note":"Angiogenin"},{"commentType":"WEB RESOURCE","resourceName":"Atlas of Genetics and Cytogenetics in Oncology and Haematology","resourceUrl":"https://atlasgeneticsoncology.org/gene/635/ANG","ftp":false}],"features":[{"type":"Signal","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":24,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2866794"}]},{"type":"Chain","location":{"start":{"value":25,"modifier":"EXACT"},"end":{"value":147,"modifier":"EXACT"}},"description":"Angiogenin","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2866794"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2866795"}],"featureId":"PRO_0000030843"},{"type":"Motif","location":{"start":{"value":55,"modifier":"EXACT"},"end":{"value":59,"modifier":"EXACT"}},"description":"Nucleolar localization signal","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"7945327"}]},{"type":"Active site","location":{"start":{"value":37,"modifier":"EXACT"},"end":{"value":37,"modifier":"EXACT"}},"description":"Proton 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