{"results":[{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"P05093","secondaryAccessions":["Q5TZV7"],"uniProtkbId":"CP17A_HUMAN","entryAudit":{"firstPublicDate":"1987-08-13","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"1987-08-13","entryVersion":245,"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":"3025870"}],"value":"Steroid 17-alpha-hydroxylase/17,20 lyase"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22266943"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25301938"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27339894"}],"value":"1.14.14.19"}]},"alternativeNames":[{"fullName":{"value":"17-alpha-hydroxyprogesterone aldolase"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22266943"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25301938"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27339894"}],"value":"1.14.14.32"}]},{"fullName":{"value":"CYPXVII"}},{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"27339894"}],"value":"Cytochrome P450 17A1"}},{"fullName":{"value":"Cytochrome P450-C17"},"shortNames":[{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"8396144"}],"value":"Cytochrome P450c17"}]},{"fullName":{"value":"Steroid 17-alpha-monooxygenase"}}]},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"19793597"},{"evidenceCode":"ECO:0000312","source":"HGNC","id":"HGNC:2593"}],"value":"CYP17A1"},"synonyms":[{"value":"CYP17"},{"value":"S17AH"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22266943"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25301938"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27339894"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36640554"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9452426"},{"evidenceCode":"ECO:0000305","source":"PubMed","id":"8027220"}],"value":"A cytochrome P450 monooxygenase involved in corticoid and androgen biosynthesis (PubMed:22266943, PubMed:25301938, PubMed:27339894, PubMed:9452426). Catalyzes 17-alpha hydroxylation of C21 steroids, which is common for both pathways. A second oxidative step, required only for androgen synthesis, involves an acyl-carbon cleavage. The 17-alpha hydroxy intermediates, as part of adrenal glucocorticoids biosynthesis pathway, are precursors of cortisol (Probable) (PubMed:25301938, PubMed:9452426). Hydroxylates steroid hormones, pregnenolone and progesterone to form 17-alpha hydroxy metabolites, followed by the cleavage of the C17-C20 bond to form C19 steroids, dehydroepiandrosterone (DHEA) and androstenedione (PubMed:22266943, PubMed:25301938, PubMed:27339894, PubMed:36640554, PubMed:9452426). Has 16-alpha hydroxylase activity. Catalyzes 16-alpha hydroxylation of 17-alpha hydroxy pregnenolone, followed by the cleavage of the C17-C20 bond to form 16-alpha-hydroxy DHEA (PubMed:36640554). Also 16-alpha hydroxylates androgens, relevant for estriol synthesis (PubMed:25301938, PubMed:27339894). Mechanistically, uses molecular oxygen inserting one oxygen atom into a substrate, and reducing the second into a water molecule, with two electrons provided by NADPH via cytochrome P450 reductase (CPR; NADPH-ferrihemoprotein reductase) (PubMed:22266943, PubMed:25301938, PubMed:27339894, PubMed:9452426)"}],"commentType":"FUNCTION"},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"a C21-steroid + reduced [NADPH--hemoprotein reductase] + O2 = a 17alpha-hydroxy-C21-steroid + oxidized [NADPH--hemoprotein reductase] + H2O + 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6beta,16alpha,17alpha-trihydroxyprogesterone + oxidized [NADPH--hemoprotein reductase] + H2O + H(+)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:53220"},{"database":"Rhea","id":"RHEA-COMP:11964"},{"database":"Rhea","id":"RHEA-COMP:11965"},{"database":"ChEBI","id":"CHEBI:763"},{"database":"ChEBI","id":"CHEBI:15377"},{"database":"ChEBI","id":"CHEBI:15378"},{"database":"ChEBI","id":"CHEBI:15379"},{"database":"ChEBI","id":"CHEBI:57618"},{"database":"ChEBI","id":"CHEBI:58210"},{"database":"ChEBI","id":"CHEBI:137046"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27339894"}]},"physiologicalReactions":[{"directionType":"left-to-right","reactionCrossReference":{"database":"Rhea","id":"RHEA:53221"},"evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"27339894"}]}]},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"17alpha-hydroxypregnenolone + reduced [NADPH--hemoprotein reductase] + O2 = 3beta-hydroxyandrost-5-en-17-one + acetate + oxidized [NADPH--hemoprotein reductase] + H2O + 2 H(+)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:50244"},{"database":"Rhea","id":"RHEA-COMP:11964"},{"database":"Rhea","id":"RHEA-COMP:11965"},{"database":"ChEBI","id":"CHEBI:15377"},{"database":"ChEBI","id":"CHEBI:15378"},{"database":"ChEBI","id":"CHEBI:15379"},{"database":"ChEBI","id":"CHEBI:28689"},{"database":"ChEBI","id":"CHEBI:28750"},{"database":"ChEBI","id":"CHEBI:30089"},{"database":"ChEBI","id":"CHEBI:57618"},{"database":"ChEBI","id":"CHEBI:58210"}],"ecNumber":"1.14.14.32","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22266943"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25301938"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27339894"}]},"physiologicalReactions":[{"directionType":"left-to-right","reactionCrossReference":{"database":"Rhea","id":"RHEA:50245"},"evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"22266943"}]}]},{"commentType":"CATALYTIC 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3beta,16alpha-dihydroxy-androst-5-en-17-one + oxidized [NADPH--hemoprotein reductase] + H2O + H(+)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:47220"},{"database":"Rhea","id":"RHEA-COMP:11964"},{"database":"Rhea","id":"RHEA-COMP:11965"},{"database":"ChEBI","id":"CHEBI:15377"},{"database":"ChEBI","id":"CHEBI:15378"},{"database":"ChEBI","id":"CHEBI:15379"},{"database":"ChEBI","id":"CHEBI:27771"},{"database":"ChEBI","id":"CHEBI:28689"},{"database":"ChEBI","id":"CHEBI:57618"},{"database":"ChEBI","id":"CHEBI:58210"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27339894"}]},"physiologicalReactions":[{"directionType":"left-to-right","reactionCrossReference":{"database":"Rhea","id":"RHEA:47221"},"evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"27339894"}]}]},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"androst-4-ene-3,17-dione + reduced [NADPH--hemoprotein reductase] + O2 = 16alpha-hydroxyandrost-4-ene-3,17-dione + oxidized [NADPH--hemoprotein reductase] + H2O + H(+)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:53228"},{"database":"Rhea","id":"RHEA-COMP:11964"},{"database":"Rhea","id":"RHEA-COMP:11965"},{"database":"ChEBI","id":"CHEBI:15377"},{"database":"ChEBI","id":"CHEBI:15378"},{"database":"ChEBI","id":"CHEBI:15379"},{"database":"ChEBI","id":"CHEBI:16422"},{"database":"ChEBI","id":"CHEBI:27582"},{"database":"ChEBI","id":"CHEBI:57618"},{"database":"ChEBI","id":"CHEBI:58210"}],"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27339894"}]},"physiologicalReactions":[{"directionType":"left-to-right","reactionCrossReference":{"database":"Rhea","id":"RHEA:53229"},"evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"27339894"}]}]},{"commentType":"COFACTOR","cofactors":[{"name":"heme","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22266943"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25301938"}],"cofactorCrossReference":{"database":"ChEBI","id":"CHEBI:30413"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10720067"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27339894"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9452426"}],"value":"Regulated predominantly by intracellular cAMP levels (PubMed:10720067). The 17,20-lyase activity is stimulated by cytochrome b5, which acts as an allosteric effector increasing the Vmax of the lyase activity (PubMed:27339894, PubMed:9452426)"}],"commentType":"ACTIVITY REGULATION"},{"commentType":"BIOPHYSICOCHEMICAL PROPERTIES","kineticParameters":{"michaelisConstants":[{"constant":10.5,"unit":"uM","substrate":"progesterone (17-alpha hydroxylation)","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25301938"}]},{"constant":5.87,"unit":"uM","substrate":"progesterone (17-alpha hydroxylation)","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36640554"}]},{"constant":0.93,"unit":"uM","substrate":"pregnenolone (17-alpha hydroxylation)","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25301938"}]},{"constant":1.19,"unit":"uM","substrate":"pregnenolone (17-alpha hydroxylation)","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36640554"}]},{"constant":1.2,"unit":"uM","substrate":"17alpha-hydroxypregnenolone (17,20 lyase activity)","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25301938"}]},{"constant":21.9,"unit":"uM","substrate":"17alpha-hydroxyprogesterone (17,20 lyase activity)","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36640554"}]}],"note":{"texts":[{"value":"kcat is 1.01 min(-1) with progesterone as substrate. kcat is 0.39 min(-1) with pregnenolone as substrate. kcat is 0.24 min(-1) with 17alpha-hydroxypregnenolone as substrate."}]}}},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22266943"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25301938"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"27339894"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9452426"}],"value":"Steroid hormone biosynthesis"}],"commentType":"PATHWAY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25301938"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9452426"}],"value":"Steroid biosynthesis; glucocorticoid biosynthesis"}],"commentType":"PATHWAY"},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"2808364"}],"value":"Endoplasmic reticulum membrane","id":"SL-0097"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"2808364"}],"value":"Microsome membrane","id":"SL-0165"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10720067"}],"value":"Phosphorylation is necessary for 17,20-lyase, but not for 17-alpha-hydroxylase activity"}],"commentType":"PTM"},{"commentType":"DISEASE","disease":{"diseaseId":"Adrenal hyperplasia 5","diseaseAccession":"DI-00045","acronym":"AH5","description":"A form of congenital adrenal hyperplasia, a common recessive disease due to defective synthesis of cortisol. Congenital adrenal hyperplasia is characterized by androgen excess leading to ambiguous genitalia in affected females, rapid somatic growth during childhood in both sexes with premature closure of the epiphyses and short adult stature. Four clinical types: 'salt wasting' (SW, the most severe type), 'simple virilizing' (SV, less severely affected patients), with normal aldosterone biosynthesis, 'non-classic form' or late-onset (NC or LOAH) and 'cryptic' (asymptomatic).","diseaseCrossReference":{"database":"MIM","id":"202110"},"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10720067"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11549685"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11836339"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12466376"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"14671162"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"1515452"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"1714904"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"1740503"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19793597"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"24140098"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"24498484"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25650406"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2808364"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8027220"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8245018"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8345056"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8396144"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8550762"},{"evidenceCode":"ECO:0000269","source":"Reference","id":"Ref.24"}]},"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 cytochrome P450 family"}],"commentType":"SIMILARITY"}],"features":[{"type":"Chain","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":508,"modifier":"EXACT"}},"description":"Steroid 17-alpha-hydroxylase/17,20 lyase","featureId":"PRO_0000051931"},{"type":"Binding site","location":{"start":{"value":202,"modifier":"EXACT"},"end":{"value":202,"modifier":"EXACT"}},"description":"","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25301938"}],"ligand":{"name":"substrate"}},{"type":"Binding site","location":{"start":{"value":442,"modifier":"EXACT"},"end":{"value":442,"modifier":"EXACT"}},"description":"axial binding 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