HSD17B4

UniProt ID: P51659
Organism: Homo sapiens
Review Status: INITIALIZED
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Gene Description

HSD17B4 encodes the peroxisomal multifunctional enzyme type 2 (MFE-2/MFP-2), also known as D-bifunctional protein (DBP), a 736-residue enzyme of the peroxisomal matrix that catalyzes the second (hydration) and third (dehydrogenation) reactions of peroxisomal fatty-acid beta-oxidation with D (R) stereochemistry. It is a multidomain protein: an N-terminal (3R)-hydroxyacyl-CoA dehydrogenase domain of the short-chain dehydrogenase/reductase (SDR) family with an NAD(+)-binding Rossmann fold (historically named 17-beta-hydroxysteroid dehydrogenase type 4), a central D-specific enoyl-CoA hydratase 2 / (3R)-3-hydroxyacyl-CoA dehydratase domain of the MaoC/hot-dog fold, and a C-terminal SCP2-like sterol-carrier / lipid-transfer domain. The full-length protein is bifunctional and forms homodimers; in vivo it is proteolytically processed into separate dehydrogenase and hydratase chains. DBP is the main bifunctional enzyme handling very-long-chain fatty acids, 2-methyl-branched-chain (pristanic) acids, C27 bile-acid intermediates (di- and trihydroxycholestanoic acid), and long-chain dicarboxylic acids, and it participates in the peroxisomal beta-oxidation step that retroconverts C24:6n-3 to docosahexaenoic acid (DHA). Its 3-oxoacyl-CoA product is passed to the peroxisomal thiolase step (SCPx/ACAA1). Loss of function causes D-bifunctional protein deficiency, a severe Zellweger-like neonatal peroxisomal disorder, while milder hypomorphic alleles cause Perrault syndrome (sensorineural deafness with ovarian dysgenesis).

Existing Annotations Review

GO Term Evidence Action Reason
GO:0106386 (3R)-3-hydroxyacyl-CoA dehydrogenase (NAD+) activity
IBA
GO_REF:0000033
ACCEPT
Summary: Phylogenetically-inferred (3R)-3-hydroxyacyl-CoA dehydrogenase (NAD+) activity, the N-terminal dehydrogenase catalytic activity of DBP. This is one of the two core enzymatic functions and is directly supported by biochemistry (see IDA annotations to the same term).
Supporting Evidence:
file:human/HSD17B4/HSD17B4-uniprot.txt
a (3R)-3-hydroxyacyl-CoA + NAD(+) = a 3-oxoacyl-CoA + NADH +
GO:0005777 peroxisome
IBA
GO_REF:0000033
ACCEPT
Summary: Phylogenetically-inferred peroxisomal localization; correct. DBP is a peroxisomal-matrix enzyme with a C-terminal PTS1-like targeting motif, consistent with IDA immunofluorescence.
Supporting Evidence:
file:human/HSD17B4/HSD17B4-uniprot.txt
SUBCELLULAR LOCATION: Peroxisome
GO:0006635 fatty acid beta-oxidation
IBA
GO_REF:0000033
ACCEPT
Summary: Phylogenetically-inferred involvement in fatty acid beta-oxidation, the core biological process for DBP (it catalyzes reactions 2 and 3 of the peroxisomal beta-oxidation cycle).
Supporting Evidence:
file:human/HSD17B4/HSD17B4-uniprot.txt
Bifunctional enzyme acting on the peroxisomal fatty acid
GO:0005777 peroxisome
IEA
GO_REF:0000120
ACCEPT
Summary: Electronic peroxisome localization (UniProt subcellular-location/ARBA). Correct and redundant with the experimentally-supported peroxisome/peroxisomal-matrix annotations.
GO:0006629 lipid metabolic process
IEA
GO_REF:0000117
MARK AS OVER ANNOTATED
Summary: Electronic ARBA annotation to the very general parent term 'lipid metabolic process'. Not wrong but far less informative than fatty acid beta-oxidation; a high-level grouping term.
GO:0016616 oxidoreductase activity, acting on the CH-OH group of donors, NAD or NADP as acceptor
IEA
GO_REF:0000117
MODIFY
Summary: Electronic ARBA annotation to a broad oxidoreductase grouping term. This is a correct but non-specific ancestor of the actual (3R)-3-hydroxyacyl-CoA dehydrogenase (NAD+) activity; the specific term GO:0106386 should be used instead.
GO:0018812 3-hydroxyacyl-CoA dehydratase activity
IEA
GO_REF:0000120
MODIFY
Summary: Electronic annotation to the dehydratase activity of the central hydratase-2 domain. Correct in essence; the D-specific term GO:0080023 ((2E)-enoyl-CoA hydratase activity, i.e. (3R)-3-hydroxyacyl-CoA = (2E)-enoyl-CoA + H2O) is the stereochemistry-precise term.
GO:0033989 3alpha,7alpha,12alpha-trihydroxy-5beta-cholest-24-enoyl-CoA hydratase activity
IEA
GO_REF:0000120
KEEP AS NON CORE
Summary: Electronic Rhea/EC-based annotation for the bile-acid (THCA-CoA) hydratase reaction (EC 4.2.1.107), a substrate-specific instance of DBP's hydratase-2 activity. This is a genuine, experimentally-supported substrate (di/trihydroxycholestanoic acid) so the term is retained as a substrate-specific non-core activity of the central hydratase domain.
Supporting Evidence:
file:human/HSD17B4/HSD17B4-uniprot.txt
EC=4.2.1.107
GO:0080023 (2E)-enoyl-CoA hydratase activity
IEA
GO_REF:0000120
ACCEPT
Summary: Electronic Rhea-based annotation to the D-specific (2E)-enoyl-CoA hydratase activity (EC 4.2.1.119); reaction (3R)-3-hydroxyacyl-CoA = (2E)-enoyl-CoA + H2O. This is the precise term for DBP's central hydratase-2 catalytic activity and is one of the two core functions.
Supporting Evidence:
file:human/HSD17B4/HSD17B4-uniprot.txt
EC=4.2.1.119
GO:0102158 very-long-chain (3R)-3-hydroxyacyl-CoA dehydratase activity
IEA
GO_REF:0000116
KEEP AS NON CORE
Summary: Electronic Rhea-based annotation to the VLCFA-specific instance of DBP's D-hydratase activity. Consistent with DBP being the main hydratase for very-long-chain fatty acids; a substrate-specific child of the core hydratase-2 activity.
Supporting Evidence:
file:human/HSD17B4/HSD17B4-uniprot.txt
a very-long-chain (3R)-3-hydroxyacyl-CoA = a very-long-chain
GO:0106386 (3R)-3-hydroxyacyl-CoA dehydrogenase (NAD+) activity
IEA
GO_REF:0000116
ACCEPT
Summary: Electronic Rhea-based annotation to the core dehydrogenase activity. Correct and redundant with the IDA/IBA annotations to the same term.
Supporting Evidence:
file:human/HSD17B4/HSD17B4-uniprot.txt
a (3R)-3-hydroxyacyl-CoA + NAD(+) = a 3-oxoacyl-CoA + NADH +
GO:0006635 fatty acid beta-oxidation
IEA
GO_REF:0000041
ACCEPT
Summary: Electronic UniPathway-based annotation to fatty acid beta-oxidation, the core process. Correct and redundant with the IDA/IBA evidence.
Supporting Evidence:
file:human/HSD17B4/HSD17B4-uniprot.txt
PATHWAY: Lipid metabolism; fatty acid beta-oxidation.
GO:0005777 peroxisome
IDA
GO_REF:0000052
ACCEPT
Summary: Immunofluorescence (HPA) peroxisome localization. Directly supports the core peroxisomal localization of DBP.
GO:0106386 (3R)-3-hydroxyacyl-CoA dehydrogenase (NAD+) activity
IDA
PMID:10706581
Peroxisomal fatty acid oxidation disorders and 58 kDa sterol...
ACCEPT
Summary: Direct assay evidence for the (3R)-3-hydroxyacyl-CoA dehydrogenase activity, measured via recombinant human D-bifunctional protein used to generate the bile-acid dehydrogenation product. Core catalytic activity.
Supporting Evidence:
PMID:10706581
expressing human D-bifunctional protein
GO:0106386 (3R)-3-hydroxyacyl-CoA dehydrogenase (NAD+) activity
IDA
PMID:15060085
Identification of the peroxisomal beta-oxidation enzymes inv...
ACCEPT
Summary: Direct enzyme-activity measurement of recombinant human DBP dehydrogenase on long-chain and dicarboxylic acyl-CoA substrates. Core catalytic activity.
Supporting Evidence:
PMID:15060085
direct enzyme measurements with human recombinant
GO:0106386 (3R)-3-hydroxyacyl-CoA dehydrogenase (NAD+) activity
IDA
PMID:9089413
Structure of D-3-hydroxyacyl-CoA dehydratase/D-3-hydroxyacyl...
ACCEPT
Summary: Direct biochemical characterization of the purified/recombinant human bifunctional protein showing dehydrogenase activity; the full-length cDNA product has both dehydratase and dehydrogenase activities. Core catalytic activity, with defined kinetics.
Supporting Evidence:
PMID:9089413
exhibited both the dehydratase and dehydrogenase activities
GO:0106386 (3R)-3-hydroxyacyl-CoA dehydrogenase (NAD+) activity
IDA
PMID:9482850
Peroxisomal D-hydroxyacyl-CoA dehydrogenase deficiency: reso...
ACCEPT
Summary: Direct evidence that the D-bifunctional protein has 3-hydroxyacyl-CoA dehydrogenase activity, and expression of the Gly16Ser variant showed loss of this activity, confirming the dehydrogenase domain. Core catalytic activity.
Supporting Evidence:
PMID:9482850
enoyl-CoA hydratase and 3-hydroxyacyl-CoA dehydrogenase activity
GO:0106386 (3R)-3-hydroxyacyl-CoA dehydrogenase (NAD+) activity
IMP
PMID:9482850
Peroxisomal D-hydroxyacyl-CoA dehydrogenase deficiency: reso...
ACCEPT
Summary: Mutant-phenotype evidence (Gly16Ser in the NAD+-binding loop) abolishing the dehydrogenase activity, on the processed dehydrogenase chain (PRO_0000400082). Confirms the N-terminal dehydrogenase catalytic activity is required and mapped to this domain.
Supporting Evidence:
PMID:9482850
the G46A mutation causes the loss of the 3-hydroxyacyl-CoA dehydrogenase activity
GO:0004303 estradiol 17-beta-dehydrogenase [NAD(P)+] activity
IDA
PMID:7487879
Molecular cloning of a novel widely expressed human 80 kDa 1...
MARK AS OVER ANNOTATED
Summary: From the original cloning paper, which reported a unidirectional oxidative 17-beta-HSD (estradiol dehydrogenase) activity when the enzyme was over-expressed in mammalian cells. This is the historical basis of the 17-beta-HSD type 4 name, but the physiological substrates are fatty-acyl-CoAs, not steroids; UniProt's FUNCTION section describes only fatty-acid/bile-acid metabolism. This steroid activity is an in-vitro side reaction and an over-annotation of the enzyme's core evolved role. Retained (experimental IDA) but flagged.
Supporting Evidence:
PMID:7487879
17 beta-HSD IV enzyme displays a specific unidirectional oxidative 17 beta-HSD activity
GO:0004300 enoyl-CoA hydratase activity
IDA
PMID:10706581
Peroxisomal fatty acid oxidation disorders and 58 kDa sterol...
MODIFY
Summary: Direct evidence for enoyl-CoA hydratase activity (the enoyl-CoA of the bile-acid intermediate THCA is hydrated by recombinant human DBP). Correct; this is the parent of the D-specific hydratase-2 activity. MODIFY to the stereochemistry-precise term GO:0080023.
Supporting Evidence:
PMID:10706581
preincubating the enoyl-CoA of the bile acid intermediate THCA
GO:0004300 enoyl-CoA hydratase activity
IDA
PMID:15060085
Identification of the peroxisomal beta-oxidation enzymes inv...
MODIFY
Summary: Direct enzyme-activity measurement of the DBP hydratase on long-chain / dicarboxylic substrates. Correct but non-specific; MODIFY to the D-specific hydratase-2 term GO:0080023.
Supporting Evidence:
PMID:15060085
direct enzyme measurements with human recombinant
GO:0004300 enoyl-CoA hydratase activity
IDA
PMID:9482850
Peroxisomal D-hydroxyacyl-CoA dehydrogenase deficiency: reso...
MODIFY
Summary: Direct evidence that DBP has enoyl-CoA hydratase activity; the enoyl-CoA hydratase activity was normal in a patient with an isolated dehydrogenase defect, confirming the distinct hydratase function. MODIFY to the D-specific term GO:0080023.
Supporting Evidence:
PMID:9482850
the enoyl-CoA hydratase activity of d -bifunctional protein was normal
GO:0006635 fatty acid beta-oxidation
IDA
PMID:10706581
Peroxisomal fatty acid oxidation disorders and 58 kDa sterol...
ACCEPT
Summary: Direct evidence connecting DBP to peroxisomal beta-oxidation of bile-acid intermediates. Core biological process.
Supporting Evidence:
PMID:10706581
peroxisomal oxidation of branched-chain fatty acids
GO:0006635 fatty acid beta-oxidation
IDA
PMID:15060085
Identification of the peroxisomal beta-oxidation enzymes inv...
ACCEPT
Summary: Direct evidence that DBP is one of the main enzymes in peroxisomal beta-oxidation of long-chain dicarboxylic acids. Core biological process.
Supporting Evidence:
PMID:15060085
the main enzymes involved in beta-oxidation of C16DCA are SCOX, both LBP and DBP
GO:0006635 fatty acid beta-oxidation
IDA
PMID:9482850
Peroxisomal D-hydroxyacyl-CoA dehydrogenase deficiency: reso...
ACCEPT
Summary: Direct evidence that DBP plays an essential, non-redundant role in the peroxisomal beta-oxidation pathway. Core biological process.
Supporting Evidence:
PMID:9482850
plays an essential role in the peroxisomal Ξ²-oxidation pathway
GO:0005782 peroxisomal matrix
TAS
Reactome:R-HSA-192331
ACCEPT
Summary: Reactome TAS localizing DBP to the peroxisomal matrix in the context of the bile-acid (THCA-CoA hydration) reaction. Correct and the most precise localization term; the peroxisomal matrix is the functional compartment for DBP catalysis.
GO:0005782 peroxisomal matrix
TAS
Reactome:R-HSA-193455
ACCEPT
Summary: Reactome TAS localizing DBP to the peroxisomal matrix (bile-acid intermediate dehydrogenation reaction). Correct core localization.
GO:0005782 peroxisomal matrix
TAS
Reactome:R-HSA-193508
ACCEPT
Summary: Reactome TAS localizing DBP to the peroxisomal matrix (bile-acid intermediate reaction). Correct core localization.
GO:0005782 peroxisomal matrix
TAS
Reactome:R-HSA-193535
ACCEPT
Summary: Reactome TAS localizing DBP to the peroxisomal matrix (bile-acid intermediate hydration reaction). Correct core localization.
GO:0005782 peroxisomal matrix
TAS
Reactome:R-HSA-2066778
ACCEPT
Summary: Reactome TAS localizing DBP to the peroxisomal matrix (hydration step in DHA-synthesis beta-oxidation). Correct core localization.
GO:0005782 peroxisomal matrix
TAS
Reactome:R-HSA-2066780
ACCEPT
Summary: Reactome TAS localizing DBP to the peroxisomal matrix (dehydrogenation step in DHA-synthesis beta-oxidation). Correct core localization.
GO:0005782 peroxisomal matrix
TAS
Reactome:R-HSA-389986
ACCEPT
Summary: Reactome TAS localizing DBP to the peroxisomal matrix (pristanoyl-CoA hydration step). Correct core localization.
GO:0005782 peroxisomal matrix
TAS
Reactome:R-HSA-389995
ACCEPT
Summary: Reactome TAS localizing DBP to the peroxisomal matrix (pristanoyl-CoA dehydrogenation step). Correct core localization.
GO:0005782 peroxisomal matrix
TAS
Reactome:R-HSA-390251
ACCEPT
Summary: Reactome TAS localizing DBP to the peroxisomal matrix (VLCFA 3-hydroxyacyl-CoA dehydrogenation step). Correct core localization.
GO:0005782 peroxisomal matrix
TAS
Reactome:R-HSA-390252
ACCEPT
Summary: Reactome TAS localizing DBP to the peroxisomal matrix (VLCFA enoyl-CoA hydration step). Correct core localization.
GO:0005782 peroxisomal matrix
TAS
Reactome:R-HSA-9033235
ACCEPT
Summary: Reactome TAS localizing DBP to the peroxisomal matrix as PEX5 import cargo (translocation from cytosol to matrix). Correct destination compartment for the imported enzyme.
GO:0005829 cytosol
TAS
Reactome:R-HSA-9033235
KEEP AS NON CORE
Summary: Reactome TAS placing DBP in the cytosol as newly-synthesized PTS1 cargo before import. This reflects the transient import route, not the functional compartment; DBP acts in the peroxisomal matrix. Non-core.
GO:0005829 cytosol
TAS
Reactome:R-HSA-9033236
KEEP AS NON CORE
Summary: Reactome TAS placing DBP in the cytosol as PEX5 cargo binding the docking/translocation module. Import-route intermediate, not the functional compartment. Non-core.
GO:0001649 osteoblast differentiation
HDA
PMID:16210410
Differential expression profiling of membrane proteins by qu...
MARK AS OVER ANNOTATED
Summary: High-throughput proteomics screen of membrane-protein expression changes in a mesenchymal stem cell line undergoing osteoblast differentiation. DBP was detected as a differentially expressed protein, not shown to have any functional role in differentiation. An expression correlation, not a functional involvement; over-annotation.
Supporting Evidence:
PMID:16210410
changes in expression of membrane protein markers
GO:0016020 membrane
HDA
PMID:16210410
Differential expression profiling of membrane proteins by qu...
MARK AS OVER ANNOTATED
Summary: From a membrane-protein proteomics screen; DBP co-fractionated in a membrane-enriched preparation. DBP is a soluble peroxisomal-matrix enzyme, not an integral membrane protein; this is a co-fractionation artifact / over-annotation of location.
Supporting Evidence:
PMID:16210410
expression profiling of membrane proteins by quantitative
GO:0005777 peroxisome
IDA
PMID:15599942
Peroxisomal branched chain fatty acid beta-oxidation pathway...
ACCEPT
Summary: Direct evidence (branched-chain fatty-acid beta-oxidation study in prostate) supporting peroxisomal localization and activity of DBP. Correct core localization.
Supporting Evidence:
PMID:15599942
D-bifunctional protein (DBP)-is also upregulated in prostate cancer
GO:0042803 protein homodimerization activity
IDA
PMID:15644212
Crystal structure of 2-enoyl-CoA hydratase 2 from human pero...
KEEP AS NON CORE
Summary: Crystal structure of the human MFE-2 hydratase-2 domain, which forms a hot-dog-fold dimer; the full-length enzyme is a homodimer. Homodimerization is a real structural property but is a supporting feature of the active enzyme rather than a distinct core molecular function; retained as non-core.
Supporting Evidence:
PMID:15644212
having a two-domain subunit structure
GO:0018812 3-hydroxyacyl-CoA dehydratase activity
IDA
PMID:9482850
Peroxisomal D-hydroxyacyl-CoA dehydrogenase deficiency: reso...
MODIFY
Summary: Direct evidence for the D-hydroxyacyl-CoA dehydratase / enoyl-CoA hydratase activity of DBP (normal hydratase activity in a dehydrogenase-defect patient). Correct; MODIFY to the stereochemistry-precise D term GO:0080023.
Supporting Evidence:
PMID:9482850
the enoyl-CoA hydratase activity of d -bifunctional protein was normal
GO:0005777 peroxisome
NAS
PMID:9482850
Peroxisomal D-hydroxyacyl-CoA dehydrogenase deficiency: reso...
ACCEPT
Summary: Non-traceable author statement that DBP is a peroxisomal beta-oxidation enzyme. Correct core localization, supported more strongly by IDA/TAS annotations.
Supporting Evidence:
PMID:9482850
a new peroxisomal Ξ²-oxidation enzyme was recently described
GO:0036111 very long-chain fatty-acyl-CoA metabolic process
IDA
PMID:9482850
Peroxisomal D-hydroxyacyl-CoA dehydrogenase deficiency: reso...
KEEP AS NON CORE
Summary: Direct evidence that DBP is involved in VLCFA metabolism: DBP-deficient patients accumulate straight-chain very-long-chain fatty acids, and DBP may be the main VLCFA beta-oxidation enzyme. A biologically important substrate class; keep, though it is a specialization of the core fatty acid beta-oxidation process.
Supporting Evidence:
PMID:9482850
accumulates straight-chain fatty acids (VLCFA) both in plasma and fibroblasts
GO:0018812 3-hydroxyacyl-CoA dehydratase activity
IDA
PMID:9089413
Structure of D-3-hydroxyacyl-CoA dehydratase/D-3-hydroxyacyl...
MODIFY
Summary: Direct biochemical evidence: the purified human bifunctional protein has dehydratase activity, and a truncated C-terminal fragment retains only dehydratase activity, mapping it to the hydratase-2 domain. Correct; MODIFY to the D-specific term GO:0080023.
Supporting Evidence:
PMID:9089413
exhibited only the dehydratase activity
GO:0036112 medium-chain fatty-acyl-CoA metabolic process
IDA
PMID:9089413
Structure of D-3-hydroxyacyl-CoA dehydratase/D-3-hydroxyacyl...
KEEP AS NON CORE
Summary: Direct evidence from kinetic characterization on medium-chain substrates (e.g. D-3-hydroxy- octanoyl-CoA, decanoyl-CoA). A substrate-specific specialization of DBP's beta-oxidation activity; retained as non-core.
Supporting Evidence:
file:human/HSD17B4/HSD17B4-uniprot.txt
KM=10 uM for D-3-hydroxy-octanoyl-CoA
GO:0042803 protein homodimerization activity
IDA
PMID:9089413
Structure of D-3-hydroxyacyl-CoA dehydratase/D-3-hydroxyacyl...
KEEP AS NON CORE
Summary: Direct evidence that the native enzyme is a homodimer (of the 77-kDa polypeptide). Structural property supporting the active enzyme; retained as non-core.
Supporting Evidence:
PMID:9089413
the native enzyme is a homodimer of 77-kDa
GO:0005778 peroxisomal membrane
HDA
PMID:21525035
PEX14 is required for microtubule-based peroxisome motility ...
MARK AS OVER ANNOTATED
Summary: HDA localization to the peroxisomal membrane, from a proteomics study whose subject is PEX14 and microtubule-based peroxisome motility (DBP is not its focus). DBP is a soluble matrix enzyme; membrane co-purification reflects peroxisome-membrane proteome preparation, not integral membrane localization. Over-annotation of location.
Supporting Evidence:
PMID:21525035
isolating native peroxisomal membrane protein complexes from cultured human cells
GO:0006635 fatty acid beta-oxidation
IDA
PMID:10400999
Enoyl-CoA hydratase deficiency: identification of a new type...
ACCEPT
Summary: Direct evidence: DBP is involved in peroxisomal beta-oxidation of VLCFAs, branched-chain fatty acids and bile-acid intermediates; a new hydratase-domain deficiency was defined here. Core biological process.
Supporting Evidence:
PMID:10400999
D-bifunctional protein is involved in the peroxisomal beta-oxidation of very long chain fatty acids, branched chain fatty acids and bile acid intermediates
GO:0018812 3-hydroxyacyl-CoA dehydratase activity
IDA
PMID:10400999
Enoyl-CoA hydratase deficiency: identification of a new type...
MODIFY
Summary: Direct evidence for the enoyl-CoA hydratase (dehydratase) domain activity; two patients had an isolated defect in the enoyl-CoA hydratase domain of DBP (Asn457Tyr). Correct; MODIFY to the D-specific term GO:0080023.
Supporting Evidence:
PMID:10400999
an isolated defect in the enoyl-CoA hydratase domain of D-bifunctional protein
GO:0005777 peroxisome
NAS
PMID:7487879
Molecular cloning of a novel widely expressed human 80 kDa 1...
ACCEPT
Summary: Non-traceable author statement of peroxisomal localization from the original cloning paper (the C-terminal region is similar to sterol carrier protein 2, a peroxisomal marker). Correct core localization, supported more strongly by IDA/TAS annotations.
Supporting Evidence:
PMID:7487879
amino acids 596-736 which are similar to human sterol
GO:0008209 androgen metabolic process
IDA
PMID:7487879
Molecular cloning of a novel widely expressed human 80 kDa 1...
MARK AS OVER ANNOTATED
Summary: From the original cloning paper reporting 17-beta-HSD (steroid) activity in over-expressing cells. DBP's physiological substrates are fatty-acyl-CoAs, not androgens; UniProt's FUNCTION section describes only fatty-acid/bile-acid metabolism. This steroid-metabolism process is a side reaction and an over-annotation of the enzyme's core role. Retained (experimental IDA) but flagged.
Supporting Evidence:
PMID:7487879
Reactions of oestrogens and androgens at position C-17 are catalysed by 17 beta-hydroxysteroid dehydrogenases
GO:0008210 estrogen metabolic process
IDA
PMID:7487879
Molecular cloning of a novel widely expressed human 80 kDa 1...
MARK AS OVER ANNOTATED
Summary: From the original cloning paper reporting oxidative 17-beta-estradiol dehydrogenase activity in over-expressing cells. As with androgen metabolism, this is an in-vitro steroid side activity, not the enzyme's physiological (fatty-acid) role; over-annotation. Retained (experimental IDA) but flagged.
Supporting Evidence:
PMID:7487879
17 beta-HSD IV enzyme displays a specific unidirectional oxidative 17 beta-HSD activity

Core Functions

D-specific enoyl-CoA hydratase 2 (D-3-hydroxyacyl-CoA dehydratase) activity: the central MaoC/hot-dog-fold domain hydrates a (2E)-enoyl-CoA to a (3R)-3-hydroxyacyl-CoA (EC 4.2.1.119), the second reaction of each peroxisomal beta-oxidation cycle, acting on very-long-chain, 2-methyl-branched (pristanic) and bile-acid-intermediate acyl-CoAs in the peroxisomal matrix.

Directly Involved In:
Cellular Locations:
Supporting Evidence:
  • file:human/HSD17B4/HSD17B4-uniprot.txt
    hydration of 2-enoyl-CoA (trans-2-enoyl-CoA) to
  • PMID:9089413
    exhibited only the dehydratase activity

(3R)-3-hydroxyacyl-CoA dehydrogenase (NAD+) activity: the N-terminal SDR/Rossmann-fold domain oxidizes a (3R)-3-hydroxyacyl-CoA with NAD+ to a 3-oxoacyl-CoA (EC 1.1.1.n12), the third reaction of each peroxisomal beta-oxidation cycle; the 3-oxoacyl-CoA product is handed to the peroxisomal thiolase step. Acts on the same VLCFA, branched-chain and bile-acid substrates.

Supporting Evidence:
  • file:human/HSD17B4/HSD17B4-uniprot.txt
    dehydrogenation of (3R)-3-
  • PMID:9482850
    enoyl-CoA hydratase and 3-hydroxyacyl-CoA dehydrogenase activity

References

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Suggested Questions for Experts

Q: Given that DBP's physiological substrates are fatty-acyl-CoAs, should the historical 17-beta-hydroxysteroid dehydrogenase (estradiol/androgen/estrogen) activities be retained as curated functions, or reclassified as in-vitro side activities?

Q: Which peroxisomal beta-oxidation substrate classes (VLCFA vs pristanic/bile-acid vs dicarboxylic vs PUFA/DHA) depend most non-redundantly on DBP versus the L-bifunctional protein (EHHADH)?

Suggested Experiments

Experiment: Substrate-resolved metabolomics of DBP-null versus EHHADH-null cells to quantify the non-redundant contribution of DBP to each acyl-CoA class (VLCFA, pristanoyl, DHCA/THCA, C16DCA, C24:6n-3).

Experiment: Structure-guided mutagenesis of the hydratase-2 substrate-binding loops to test whether the reported steroid dehydrogenase activity and the fatty-acyl dehydrogenase activity can be separated, clarifying the physiological relevance of the 17-beta-HSD activity.

πŸ“š Additional Documentation

Notes

(HSD17B4-notes.md)

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