Information on EC 1.3.1.72 - DELTA24-sterol reductase

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The enzyme appears in viruses and cellular organisms

EC NUMBER
COMMENTARY hide
1.3.1.72
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RECOMMENDED NAME
GeneOntology No.
DELTA24-sterol reductase
REACTION
REACTION DIAGRAM
COMMENTARY hide
ORGANISM
UNIPROT
LITERATURE
5alpha-cholest-7-en-3beta-ol + NADP+ = 5alpha-cholesta-7,24-dien-3beta-ol + NADPH + H+
show the reaction diagram
REACTION TYPE
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
oxidation
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-
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redox reaction
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reduction
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PATHWAY
BRENDA Link
KEGG Link
MetaCyc Link
cholesterol biosynthesis I
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cholesterol biosynthesis II (via 24,25-dihydrolanosterol)
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cholesterol biosynthesis III (via desmosterol)
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cholesterol biosynthesis
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Steroid biosynthesis
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Metabolic pathways
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Biosynthesis of secondary metabolites
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SYSTEMATIC NAME
IUBMB Comments
sterol:NADP+ DELTA24-oxidoreductase
Acts on a range of steroids with a 24(25)-double bond, including lanosterol, desmosterol and zymosterol.
CAS REGISTRY NUMBER
COMMENTARY hide
9033-57-2
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ORGANISM
COMMENTARY hide
LITERATURE
UNIPROT
SEQUENCE DB
SOURCE
var. atropurpurea
A0A0P0UTN5
UniProt
Manually annotated by BRENDA team
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-
-
Manually annotated by BRENDA team
tobacco hornworm
-
-
Manually annotated by BRENDA team
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-
-
Manually annotated by BRENDA team
hybrid Nongda 3138
SwissProt
Manually annotated by BRENDA team
GENERAL INFORMATION
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
malfunction
metabolism
physiological function
additional information
SUBSTRATE
PRODUCT                       
REACTION DIAGRAM
ORGANISM
UNIPROT
COMMENTARY
(Substrate) hide
LITERATURE
(Substrate)
COMMENTARY
(Product) hide
LITERATURE
(Product)
Reversibility
r=reversible
ir=irreversible
?=not specified
24-methylenecholesterol + NADPH + H+
campesterol + dihydrobrassicasterol + NADP+
show the reaction diagram
-
-
-
-
?
24-methylenecholesterol + NADPH + H+
campesterol + NADP+
show the reaction diagram
4,4-dimethyl-5alpha-cholesta-8,14,24-trien-3beta-ol + NADPH
4,4-dimethyl-5alpha-cholesta-8,14-dien-3beta-ol + NADP+
show the reaction diagram
-
-
-
?
4,4-dimethyl-5alpha-cholesta-8,24-dien-3beta-ol + NADPH
4,4-dimethyl-5alpha-cholesta-8-en-3beta-ol + NADP+
show the reaction diagram
5alpha-cholest-7-en-3beta-ol + NADP+
5alpha-cholesta-7,24-dien-3beta-ol + NADPH + H+
show the reaction diagram
-
-
-
-
r
5alpha-cholesta-7,24-dien-3beta-ol + NADPH
5alpha-cholest-7-en-3beta-ol + NADP+
show the reaction diagram
5alpha-cholesta-7,24-dien-3beta-ol + NADPH
5alpha-cholesta-7-en-3beta-ol + NADP+
show the reaction diagram
7-dehydrodesmosterol + NADPH
7-dehydrocholesterol + NADP+
show the reaction diagram
-
-
-
?
cholesta-5,24-dien-3beta-ol + NADPH + H+
5alpha-cholest-5-en-3beta-ol + NADP+
show the reaction diagram
cholesta-5,7,24-trien-3beta-ol + NADPH
cholest-5,7-dien-3beta-ol + NADP+
show the reaction diagram
cholesta-5,7,24-trien-3beta-ol + NADPH + H+
cholesta-5,7-dien-3beta-ol + NADP+
show the reaction diagram
-
-
-
-
?
cycloartenol + NADPH + H+
cycloartanol + NADP+
show the reaction diagram
-
-
-
?
DELTA5,7-avenasterol + NADPH + H+
DELTA5,7-sitosterol + NADP+
show the reaction diagram
DELTA7-avenasterol + NADPH + H+
DELTA7-sitosterol + NADP+
show the reaction diagram
desmosterol + FADH2
cholesterol + FAD + H+
show the reaction diagram
-
-
-
?
desmosterol + NADPH
cholesterol + NADP+
show the reaction diagram
ergosta-5,7,22,24(28)-tetraen-3beta-ol + NADPH + H+
ergosterol + NADP+
show the reaction diagram
isofucosterol + NADPH + H+
sitosterol + NADP+
show the reaction diagram
lanosterol + NADPH
24-dihydrolanosterol + NADP+
show the reaction diagram
lanosterol + NADPH
4,4,14alpha-trimethyl-5alpha-cholesta-8-en-3beta-ol + NADP+
show the reaction diagram
lanosterol + NADPH + H+
24-dihydrolanosterol + NADP+
show the reaction diagram
-
-
-
?
zymosterol + NADPH
5alpha-cholesta-8-en-3beta-ol + NADP+
show the reaction diagram
additional information
?
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NATURAL SUBSTRATES
NATURAL PRODUCTS
REACTION DIAGRAM
ORGANISM
UNIPROT
COMMENTARY
(Substrate) hide
LITERATURE
(Substrate)
COMMENTARY
(Product) hide
LITERATURE
(Product)
REVERSIBILITY
r=reversible
ir=irreversible
?=not specified
24-methylenecholesterol + NADPH + H+
campesterol + NADP+
show the reaction diagram
5alpha-cholest-7-en-3beta-ol + NADP+
5alpha-cholesta-7,24-dien-3beta-ol + NADPH + H+
show the reaction diagram
-
-
-
-
r
5alpha-cholesta-7,24-dien-3beta-ol + NADPH
5alpha-cholest-7-en-3beta-ol + NADP+
show the reaction diagram
5alpha-cholesta-7,24-dien-3beta-ol + NADPH
5alpha-cholesta-7-en-3beta-ol + NADP+
show the reaction diagram
cholesta-5,24-dien-3beta-ol + NADPH + H+
5alpha-cholest-5-en-3beta-ol + NADP+
show the reaction diagram
cholesta-5,7,24-trien-3beta-ol + NADPH
cholest-5,7-dien-3beta-ol + NADP+
show the reaction diagram
desmosterol + NADPH
cholesterol + NADP+
show the reaction diagram
lanosterol + NADPH
24-dihydrolanosterol + NADP+
show the reaction diagram
additional information
?
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COFACTOR
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
INHIBITORS
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
(2S)-2-((3S,5S,10S,13R,17R)-3-hydroxy-10,13-dimethyl-2,3,4,5,6,9,10,11,12,13,14,15,16,17-tetradecahydro-1H-cyclopenta[a]phenanthren-17-yl)-N-methylpropanamide
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1 microM, 58% inhibition of total cholesterol production. Acts both on sterol reductoase and lathosterol oxidase
(2S)-2-((3S,5S,10S,13R,17R)-3-hydroxy-10,13-dimethyl-2,3,4,5,6,9,10,11,12,13,14,15,16,17-tetradecahydro-1H-cyclopenta[a]phenanthren-17-yl)-N-propylpropanamide
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1 microM, 45% inhibition of total cholesterol production. Acts both on sterol reductoase and lathosterol oxidase
(2S)-2-((3S,5S,10S,13R,17R)-3-hydroxy-10,13-dimethyl-2,3,4,5,6,9,10,11,12,13,14,15,16,17-tetradecahydro-1H-cyclopenta[a]phenanthren-17-yl)propanamide
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1 microM, 75% inhibition of total cholesterol production
(2S)-N-ethyl-2-((3S,5S,10S,13R,17R)-3-hydroxy-10,13-dimethyl-2,3,4,5,6,9,10,11,12,13,14,15,16,17-tetradecahydro-1H-cyclopenta[a]phenanthren-17-yl)propanamide
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1 microM, 61% inhibition of total cholesterol production
(3S,5S,10S,13R,17R)-10,13-dimethyl-17-((S)-1-(methylamino)-1-oxopropan-2-yl)-2,3,4,5,6,9,10,11,12,13,14,15,16,17-tetradecahydro-1H-cyclopenta[a]phenanthren-3-yl acetate
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1 microM, 84% inhibition of total cholesterol production. Acts both on sterol reductoase and lathosterol oxidase
(3S,5S,10S,13R,17R)-10,13-dimethyl-17-((S)-1-oxo-1-(propylamino)propan-2-yl)-2,3,4,5,6,9,10,11,12,13,14,15,16,17-tetradecahydro-1H-cyclopenta[a]phenanthren-3-yl acetate
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1 microM, 71% inhibition of total cholesterol production. Acts both on sterol reductoase and lathosterol oxidase
(3S,5S,10S,13R,17R)-17-((S)-1-(ethylamino)-1-oxopropan-2-yl)-10,13-dimethyl-2,3,4,5,6,9,10,11,12,13,14,15,16,17-tetradecahydro-1H-cyclopenta[a]phenanthren-3-yl acetate
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1 microM, 97% inhibition of total cholesterol production
(3S,5S,10S,13R,17R)-17-((S)-1-amino-1-oxopropan-2-yl)-10,13-dimethyl-2,3,4,5,6,9,10,11,12,13,14,15,16,17-tetradecahydro-1H-cyclopenta[a]phenanthren-3-yl acetate
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1 microM, 94% inhibition of total cholesterol production
24(R,S),25-epimino-lanosterol
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iminiolanosterol, IL, potent inhibitor, IC50: 0.002 mM
24(S),25-epoxycholesterol
25-Azacholesterol
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3-beta-(2-(diethylamino)ethoxy)androst-5-en-17-one
5,22-cholestedien-3beta-ol
azasteroids
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mechanism of inhibition
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brassicasterol
Calmodulin antagonists
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ergosterol
N-(6-aminohexyl)-5-chloro-1-naphthalene sulfonamide
stigmasterol
tamoxifen
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inhibition mechanism and pattern, uncompetitive, 50% inhibition at 0.004 mM, cytotoxic in vivo, inhibitory effect on th whole cholesterol biosynthetic pathway, overview
Trifluoperazine
Triparanol
U18666A
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an inhibitor of DHCR24 activity, prevents ACTH-induced translocation of the enzyme to the nucleus in adrenal cells but not in prostate cancer cells
additional information
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ACTIVATING COMPOUND
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
cholesterol
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high levels of DHCR24 are associated with elevated cholesterol concentrations
cholestyramine
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120fold induction and activation by feeding 5% cholestyramine plus 0.1% lovastatin, CL-diet, and by modulating diurnal variation
CO
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substrate lanosterol: required to inhibit 14alpha-methyl demethylase, 14alpha-demethylation of lanosterol. 14alpha-methyl demethylase activity is dominant over 24-reductase activity, and blockade or removal of 14alpha-methyl demethylase activity is absolutely required for the detection of maximal 24-reductase activity when lanosterol substrate is present. 24-reductase activity is activated in time-dependent manner when 14alpha-methyl demethylase is blocked by CO treatment
FAD
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activate 2fold when added to the assay
H2O2
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DHCR24 mRNA levels increase in response to oxidative stress (0.1 mM H2O2) in a time-dependent manner, reaching the maximum after 4 h
ketoconazole
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substrate lanosterol: dose-dependent activation, less effective than miconazole, required to inhibit 14alpha-methyl demethylase, 14alpha-demethylation of lanosterol. 14alpha-methyl demethylase activity is dominant over 24-reductase activity, and blockade or removal of 14alpha-methyl demethylase activity is absolutely required for the detection of maximal 24-reductase activity when lanosterol substrate is present
lovastatin
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120fold induction and activation by feeding 5% cholestyramine plus 0.1% lovastatin, CL-diet, and by modulating diurnal variation
miconazole
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substrate lanosterol: dose-dependent activation, maximum activation with about 0.01 mM miconazole, required to inhibit 14alpha-methyl demethylase, 14alpha-demethylation of lanosterol. 14alpha-methyl demethylase activity is dominant over 24-reductase activity, and blockade or removal of 14alpha-methyl demethylase activity is absolutely required for the detection of maximal 24-reductase activity when lanosterol substrate is present
KM VALUE [mM]
SUBSTRATE
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
0.018 - 0.109
4,4',14alpha-trimethyl-5alpha-cholesta-8,24-dien-3beta-ol
0.0026 - 0.163
5alpha-cholesta-5,24-dien-3beta-ol
0.037
5alpha-cholesta-7,24-dien-3beta-ol
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0.176
5alpha-cholesta-8,24-dien-3beta-ol
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zymosterol
0.033
Cycloartenol
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26.3
desmosterol
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pH 7.6, 37°C
additional information
additional information
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Ki VALUE [mM]
INHIBITOR
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
0.000157
3-beta-(2-(diethylamino)ethoxy)androst-5-en-17-one
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0.0033
5,22-cholestedien-3beta-ol
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pH 7.6, 37°C
0.043
brassicasterol
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pH 7.6, 37°C
0.037
ergosterol
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pH 7.6, 37°C
0.041
stigmasterol
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pH 7.6, 37°C
0.0041
tamoxifen
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0.000523
Triparanol
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IC50 VALUE [mM]
INHIBITOR
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
0.002
24(R,S),25-epimino-lanosterol
Manduca sexta
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iminiolanosterol, IL, potent inhibitor, IC50: 0.002 mM
0.00015
3-beta-(2-(diethylamino)ethoxy)androst-5-en-17-one
Rattus norvegicus
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U18666A, non-competitive inhibition, Ki: 0.000157 mM, IC50 about 0.00015 mM, 690fold higher affinity for the enzyme than substrate lanosterol
0.0008
Triparanol
Rattus norvegicus
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non-competitive inhibition, specific inhibitor, Ki: 0.000523 mM, IC50 about 0.0008 mM, 208fold higher affinity for the enzyme than substrate lanosterol
SPECIFIC ACTIVITY [µmol/min/mg]
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
0.0015
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substrate: 5alpha-cholesta-7,24-dien-3beta-ol
additional information
pH OPTIMUM
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
6.2
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assay at, anaerobically
7.2
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assay at
TEMPERATURE OPTIMUM
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
SOURCE TISSUE
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
SOURCE
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intestinal mucosa cells
Manually annotated by BRENDA team
abundantly expressed throughout the epidermis except for the stratum corneum and in hair follicles
Manually annotated by BRENDA team
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fibroblast from skin
Manually annotated by BRENDA team
A0A0P0UTN5
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Manually annotated by BRENDA team
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DHCR24 is specifically expressed on the surface of HCC cell lines
Manually annotated by BRENDA team
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BE(2)C cell line
Manually annotated by BRENDA team
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a human hepatoblastoma-derived cell line
Manually annotated by BRENDA team
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human skin fibroblast
Manually annotated by BRENDA team
additional information
LOCALIZATION
ORGANISM
UNIPROT
COMMENTARY hide
GeneOntology No.
LITERATURE
SOURCE
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hepatocellular carcinoma cells, hepatoblastoma cell line, and cervical adenocarcinoma-derived cell line
Manually annotated by BRENDA team
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a predicted transmembrane (TM) domain-deleted DHCR24 mutation is localized to the cytoplasm
Manually annotated by BRENDA team
additional information
MOLECULAR WEIGHT
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
60000
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SDS-PAGE
60100
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x * 60100, sequence calculation
SUBUNITS
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
additional information
enzyme secondary structure
POSTTRANSLATIONAL MODIFICATION
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
phosphoprotein
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protein kinase C (PKC) activates DHCR24 activity through reversible phosphorylation. PKC inhibitors, BIM and Ro-318220, reduce cholesterol levels and accumulate desmosterol within 4 h, indicating decreased DHCR24 activity. Phosphorylation at T110 modulates DHCR24 activity
additional information
OXIDATION STABILITY
ORGANISM
UNIPROT
LITERATURE
seladin-1 expression is up-regulated in an acute response and down-regulated in a chronic response to oxidative stress
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688975
Purification/COMMENTARY
ORGANISM
UNIPROT
LITERATURE
partially from transgenic CHO cells by subcellular fractionation and membrane isolation
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Cloned/COMMENTARY
ORGANISM
UNIPROT
LITERATURE
construction of a recombinant adenovirus driving DHCR24 expression vector specific for neurons with neuron-specific promoter, synapsin-1 (SYN1), cloning of a SYN1 promoter DNA fragment of human enzyme DHCR24, and transfection of recombinant Ad-hSYN1-DHCR24 into HEK AD-293, N2A (mouse neuroblastoma), and MIN6 (mouse pancreatic carcinoma) cells. DHCR24 is specially expressed in HEK AD-293 and N2A cells, but not in MIN6 cells. Adenovirus transfection inhibits apoptosis through scavenging excess reactive oxygen species, and recombinant DHCR24 adenoviruse induces neuron-specific DHCR24 expression
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construction of a recombinant adenovirus driving DHCR24 expression vector specific for neurons with neuron-specific promoter, synapsin-1 (SYN1), cloning of a SYN1 promoter DNA fragment of rat enzyme DHCR24, and transfection of recombinant Ad-rSYN1-DHCR24 into HEK AD-293, N2A (mouse neuroblastoma), and MIN6 (mouse pancreatic carcinoma) cells. DHCR24 is specially expressed in HEK AD-293 and N2A cells, but not in MIN6 cells. Adenovirus transfection inhibits apoptosis through scavenging excess reactive oxygen species, and recombinant DHCR24 adenoviruse induces neuron-specific DHCR24 expression
DWF1, phylogenetic tree
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ectopic expression of wild-type and mutant human DHCR24s in CHO-7 cells that are deficient for DHCR24 through specific siRNA knockout, quantitative real-time RT-PCR enzyme expression analysis
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expressed as a EGFP-fusion protein in murine neuroblastoma cell line N2A
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expressed in CHO cells
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expressed in Mus musculus embryonic fibroblasts and PC-12 cells infected with adenovirus
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gene ArDWF1, cloning from hairy root expression sequence tag library, DNA and amino acid sequence determination and analysis, sequence comparison and phylogenetic tree, functional recombinant expression in Saccharomyces cerevisiae, ArDWF1-expressing yeast T21 cells produce 24-methylcholesterol in vivo
A0A0P0UTN5
gene DHCR24, DNA and amino acid sequence determination and analysis of wild-type and natural desmosterolysis mutant genes, chromosomal mapping to 1p31.1-p33, functional expression of wild-type and mutant enzymes in Saccharomyces cerevisiae
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gene OsDWF1, sequence comparison and phylogenetic tree, functional recombinant expression in Saccharomyces cerevisiae, OsDWF1-expressing yeast T21 cells produce 24-methylcholesterol in vivo
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gene ZmDWF1, single copy gene, DNA and amino acid sequence determination and analysis, cloning of a plant expression vector for construction of transgenic plants
generation of plasmids pcDNA5-DHCR7-myc/FRT and pcDNA5-DHCR24-V5/FRT, and of DHCR24 mutant plasmid pcDNA5-DHCR24 Y471S-V5/FRT, the latter is stably recombinantly expressed in CHO-7 cells, stable overexpression of inactive DHCR24 mutant Y471S in CHO-7 cells, quantitative real-time PCR expression analysis
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overexpressed in ATDC-5 cells
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quantitative real-time PCR DHCR24 expression analysis
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recombinant expression of C-terminally YFP-tagged wild-type enzyme in Arabidopsis thaliana mutant lines deficient in the enzyme in the endoplasmic reticulum and plasma membrane
recombinant expression of truncated DHCR24 (DELTA23 DHCR24) with a V5 epitope tag at either the N-terminus (V5-DHCR24) or C-terminus (DHCR24-V5) in CHO-7 cells, and the cellular localization of DELTA23 DHCR24 is primarily to the membrane fraction like the wild-type
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RzM6-0d cells are transduced with DHCR24 lentivirus. Expression of DHCR24 from pGEM-T easy vector via in vitro translation using reticulocyte lysate. Expression of HA- or FLAG-tagged DHCR24 in RzM6 or HepG2 cells. Expression of GFP- and FLAG-tagged DHCR24 in 293FT cells
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the DHCR24 gene locus is 1p32.3, spans about 46.4 kb, and comprises eight introns and nine exons, promoter map of human DHCR24, phylogenetic tree
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EXPRESSION
ORGANISM
UNIPROT
LITERATURE
activation of constitutive androstane receptor (CAR) in cultured human hepatocytes increases mRNA levels of mouse Dhcr24 and human DHCR24. A CAR-responsive element is identified in the promoter of human DHCR24
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activation of constitutive androstane receptor (CAR) in mouse livers increases mRNA levels of mouse Dhcr24 and human DHCR24
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DHCR24 is overexpressed in hepatitis C virus-related hepatocellular carcinoma specifically induced by hepatitis C virus
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DHCR24 is transcriptionally regulated by sterols via the sterol-regulatory element-binding protein-2 transcription factor
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expression of DHCR24 is mediated by two sterol regulatory elements (SREs) in the promoter of the gene, assisted by two nearby NF-Y binding sites
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hepatitis C virus induces the DHCR24 overexpression in human hepatocytes
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histone acetylation also regulates DHCR24 expression, with a reduction in DHCR24 transcription correlating with recruitment of acetylated histones H3 and H4 to its promoter
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the enzyme is regulated by the following transcription factors/proteins: sterol regulatory element binding protein 2, nuclear factor Y (via methylation), specificity protein 1, estrogen receptor, androgen receptor, thyroid hormone receptor, constitutive androstane receptor, and pregnane X receptor. Transcriptional regulation of DHCR24, detailed overview
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the sterol-regulated region is extremely GC rich and lies within a CpG island, a region of DNA with a high G and C content and a high frequency of CpG dinucleotides, which is conserved in mammals. In addition to containing a specificity protein 1 (Sp1) site required for augmentation of DHCR24 in HCV infection, this region is also important in epigenetic regulation of DHCR24. Methylation of this region decreases DHCR24 expression, with pronounced methylation occurring in cell types with low DHCR24 levels. Sex steroids, such as estrogens and androgens, are positive regulators of DHCR24, increasing gene expression via activation of their respective nuclear receptors, estrogen receptor and androgen receptor, overview. Adrenocorticotropic hormone (ACTH) stimulates the enzyme expression in adrenal gland
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ENGINEERING
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
E480K
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site-directed mutagenesis, mutation of a C-terminal domain residue, about 50% of wild-type activity remain
N294T/K306N
R103C
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site-directed mutagenesis, mutation of a FAD binding domain residue
R94H
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site-directed mutagenesis, mutation of a FAD binding domain residue, about 20% of wild-type activity remain
T110A
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site-directed mutagenesis, mutation and inactivation of the phosphorylation site results in 60% loss of activity compared to wild-type
T110E
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site-directed mutagenesis, DHCR24 activity of the phosphomimetic T110E mutant is similar to wild-type activity
Y299F
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site-directed mutagenesis, compared with wild-type DHCR24, the mutant Y299F stable cells contain reduced DHCR24 mRNA expression while having comparable DHCR24 protein levels, the mutant enzyme activity is reduced by 40% compared to the wild-type
Y300F
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site-directed mutagenesis, compared with wild-type DHCR24, the mutant Y299F stable cells contain comparable DHCR24 mRNA expression while having reduced DHCR24 protein levels, the mutant enzyme activity is similar to the wild-type
Y321F
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site-directed mutagenesis, the mutant enzyme activity is similar to the wild-type
Y507F
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site-directed mutagenesis, the mutant enzyme activity is reduced by 60% compared to the wild-type
additional information
APPLICATION
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
agriculture
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inhibition of phytosterol metabolism to cholesterol by interfering with DELTA24-sterol reductase activity presents a unique target site that might be exploited further as a selective, biorational insect-control technology
medicine
pharmacology
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3beta-hydroxysterol DELTA24-reductase on the surface of hepatitis C virus-related hepatocellular carcinoma cells can be a target for molecular targeting therapy