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C-P4H alpha subunit (III)
-
collagen proline hydroxylase
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collagen prolyl 4-hydroxylase
collagen prolyl 4-hydroxylase (type-II)
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collagen prolyl 4-hydroxylase 1
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collagen prolyl-4-hydroxylase
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collagen prolyl-4-hydroxylase alpha subunit 2
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egg-laying abnormal-9 prolyl hydroxylase
-
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EGLN prolyl hydroxylase
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HIF prolyl hydroxylase
-
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HIF prolyl-4-hydroxylase
-
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HIF-1alpha-specific prolyl-hydroxylase
-
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hydroxylase, collagen proline
-
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hypoxia inducible factor prolyl-4-hydroxylase domain-containing protein
-
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hypoxia inducible factor-prolyl hydroxylase
-
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hypoxia-inducible factor -1alpha-type prolyl 4-hydroxylase
hypoxia-inducible factor prolyl hydroxylase
-
-
hypoxia-inducible factor-1alpha-prolyl-hydroxylase 2
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PBCV-1 P4H
Paramecium bursaria Chlorella virus-1
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peptidyl proline hydroxylase
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procollagen prolyl 4-hydroxylase
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procollagen-proline dioxygenase
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proline 4-hydroxylase
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proline hydroxylase
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proline protocollagen hydroxylase
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proline, 2-oxoglutarate dioxygenase
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proline,2-oxoglutarate 4-dioxygenase
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prolyl 4-hydroxylase A085R
-
prolyl hydroxylase domain 1
-
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prolyl hydroxylase domain containing protein
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prolyl hydroxylase domain enzyme
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prolyl hydroxylase domain protein 2
-
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prolyl-4-hydroxylase alpha I
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prolyl-4-hydroxylase alpha II
-
-
prolyl-4-hydroxylase alpha subunit 2
-
prolyl-4-hydroxylase alpha1
prolyl-4-hydroxylase-alpha1
prolyl-glycyl-peptide, 2-oxoglutarate:oxygen oxidoreductase, 4-hydroxylating
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prolylprotocollagen dioxygenase
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prolylprotocollagen hydroxylase
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protocollagen hydroxylase
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protocollagen proline 4-hydroxylase
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protocollagen proline dioxygenase
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protocollagen proline hydroxylase
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protocollagen prolyl hydroxylase
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type I proly 4-hydroxylase
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type I proyl 4-hydroxylase
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type II proyl 4-hydroxylase
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BaP4H

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C-P4H

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C-P4H alpha(III)

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collagen prolyl 4-hydroxylase

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collagen prolyl 4-hydroxylase
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collagen prolyl 4-hydroxylase
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collagen prolyl 4-hydroxylase
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collagen prolyl 4-hydroxylase
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collagen prolyl 4-hydroxylase
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collagen prolyl 4-hydroxylase
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CP4H

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hypoxia-inducible factor -1alpha-type prolyl 4-hydroxylase

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hypoxia-inducible factor -1alpha-type prolyl 4-hydroxylase
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P4H

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P4H alpha1

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P4H1

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P4ha1

P4H alpha 1 subunit
P4ha2

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PHD

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PHD1

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PHD2

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PHD3

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prolyl 4-hydroxylase

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prolyl 4-hydroxylase
Q81LZ8
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prolyl hydroxylase

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prolyl-4-hydroxylase

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prolyl-4-hydroxylase alpha1

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prolyl-4-hydroxylase alpha1
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prolyl-4-hydroxylase alpha1
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prolyl-4-hydroxylase-alpha1

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prolyl-4-hydroxylase-alpha1
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Skp1 prolyl hydroxylase

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Skp1 prolyl hydroxylase
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additional information

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the enzyme belongs to the P4H enzyme family of the 2-oxoglutarate-dependent dioxygenases
additional information
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the enzyme belongs to the 2-oxoglutarate dioxygenase family
additional information
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the enzyme belongs to the P4H enzyme family of the 2-oxoglutarate-dependent dioxygenases
additional information
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the enzyme belongs to the P4H enzyme family of the 2-oxoglutarate-dependent dioxygenases
additional information
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the enzyme belongs to the alpha-ketoglutarate-dependent iron(II) dioxygenases
additional information
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the enzyme belongs to the P4H enzyme family of the 2-oxoglutarate-dependent dioxygenases
additional information
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the enzyme belongs to the P4H enzyme family of the 2-oxoglutarate-dependent dioxygenases
additional information
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the enzyme belongs to the P4H enzyme family of the 2-oxoglutarate-dependent dioxygenases
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2

mechanism
-
procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
mechanism
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
mechanism, kinetic analysis of the reaction sequence
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
mechanism, role of cosubstrates
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
active site and reaction mechanism, overview
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
active site structure and action mode, overview
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
reaction mechanism involving ascorbate, which are not required for the hydroxylation reaction part, but for the uncoupled cedarboxylation, overview
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
reaction mechanism involving ascorbate, which not required for the hydroxylation reaction part, but for the uncoupled cedarboxylation, overview
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
reaction mechanism involving ascorbate, which not required for the hydroxylation reaction part, but for the uncoupled cedarboxylation, overview
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
reaction mechanism involving ascorbate, which not required for the hydroxylation reaction part, but for the uncoupled cedarboxylation, overview
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
reaction mechanism involving ascorbate, which not required for the hydroxylation reaction part, but for the uncoupled cedarboxylation, overview
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
reaction mechanism involving ascorbate, which not required for the hydroxylation reaction part, but for the uncoupled cedarboxylation, overview
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
reaction mechanism with Fe2+ bound in the active site by a 2-His-1-Asp motif, overview
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procollagen L-proline + 2-oxoglutarate + O2 = procollagen trans-4-hydroxy-L-proline + succinate + CO2
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(2S,4S)-4-fluoroproline + 2-oxoglutarate + O2
(2S)-4-oxoproline + fluoride
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development and evaluation of an assay method that continuously and directly detects turnover of the proline-containing substrate. The assay is based on (2S,4S)-4-fluoroproline, a proline analogue that is transformed into (2S)-4-oxoproline and inorganic fluoride by P4H using a fluoride ion-selective electrode for detection
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?
(Ala-Pro-Gly)5 + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
recombinant enzyme
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?
(Ala-Thr-Pro-Pro-Pro-Val)3 + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
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?
(ATPPPV)3 + 2-oxoglutarate + O2
trans-4-hydroxy-L-proline (ATPPPV)3 + succinate + CO2
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?
(Gly-Ala-Pro)n + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
(Gly-Pro-4Hyp)5 + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
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?
(Gly-Pro-Ala)n + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
(Gly-Pro-Pro)10 + 2-oxoglutarate + O2
?
(GPP)10 + 2-oxoglutarate + O2
?
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?
(L-Pro-L-Ala-L-Pro-L-Lys)2 + 2-oxoglutarate + O2
(L-Pro-L-Ala-L-Pro-L-Lys)2-trans-4-hydroxy-L-proline + succinate + CO2
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r
(L-Pro-L-Ala-L-Pro-L-Lys)3 + 2-oxoglutarate + O2
(L-Pro-L-Ala-L-Pro-L-Lys)3-trans-4-hydroxy-L-proline + succinate + CO2
(L-Pro-L-Glu-L-Pro-L-Pro-L-Ala)5 L-proline + 2-oxoglutarate + O2
(L-Pro-L-Glu-L-Pro-L-Pro-L-Ala)5 trans-4-hydroxy-L-proline + succinate + CO2
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?
(L-Pro-L-Pro-Gly)10 + 2-oxoglutarate + O2
(L-Pro-L-Pro-Gly)10-trans-4-hydroxy-L-proline + succinate + CO2
(L-Pro-L-Pro-Gly)10 L-proline + 2-oxoglutarate + O2
(L-Pro-L-Pro-Gly)10 trans-4-hydroxy-L-proline + succinate + CO2
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44.8% of the activity with (L-Pro-L-Glu-L-Pro-L-Pro-L-Ala)5 L-proline
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?
(L-Pro-L-Pro-Gly)2 + 2-oxoglutarate + O2
(L-Pro-L-Pro-Gly)2-trans-4-hydroxy-L-proline + succinate + CO2
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r
(L-Pro-L-Pro-Gly)5 + 2-oxoglutarate + O2
(L-Pro-L-Pro-Gly)5-trans-4-hydroxy-L-proline + succinate + CO2
(L-prolylglycyl-L-prolyl)n + 2-oxoglutarate + O2
4-hydroxyproline containing peptide
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molecular weight of the peptide substrate: about 4000
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?
(L-Ser-L-Pro)19 L-proline + 2-oxoglutarate + O2
(L-Ser-L-Pro)19 trans-4-hydroxy-L-proline + succinate + CO2
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?
(L-Ser-L-Pro-L-Ala-L-Pro-L-Pro)5 L-proline + 2-oxoglutarate + O2
(L-Ser-L-Pro-L-Ala-L-Pro-L-Pro)5 trans-4-hydroxy-L-proline + succinate + CO2
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?
(L-Ser-L-Pro-L-Glu-L-Pro-L-Pro)5 L-proline + 2-oxoglutarate + O2
(L-Ser-L-Pro-L-Glu-L-Pro-L-Pro)5 trans-4-hydroxy-L-proline + succinate + CO2
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?
(L-Ser-L-Pro-L-Lys-L-Pro-L-Pro)5 L-proline + 2-oxoglutarate + O2
(L-Ser-L-Pro-L-Lys-L-Pro-L-Pro)5 trans-4-hydroxy-L-proline + succinate + CO2
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?
(Pro-Ala-Gly)5 + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
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?
(Pro-Ala-Pro-Lys)n + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
(Pro-Glu-Pro-Pro-Ala)5 + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
(Pro-Gly-Pro)n + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
(Pro-Pro-Gly)10 + 2-oxoglutarate + O2
? + succinate + CO2
(Pro-Pro-Gly)10 + 2-oxoglutarate + O2
trans-4-hydroxy-L-proline (Pro-Pro-Gly)10 + succinate + CO2
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?
(Pro-Pro-Gly)10 + O2
?
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?
(Pro-Pro-Gly)10 L-proline + 2-oxoglutarate + O2
(Pro-Pro-Gly)10 trans-4-hydroxy-L-proline + succinate + CO2
(Pro-Pro-Gly)2 L-proline + 2-oxoglutarate + O2
(Pro-Pro-Gly)2 trans-4-hydroxy-L-proline + succinate + CO2
(Pro-Pro-Gly)n + 2-oxoglutarate + O2
(Pro-4-hydroxy-Pro-Gly)n + succinate + CO2
(Ser-Pro)5 + O2
?
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the (Ser-Pro)5 peptide is found only in molecules A and C of the four polypeptides forming the enzyme
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?
(Ser-Pro-Lys-Pro-Pro)5 + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
100-amino acid alpha1(I) collagen fragment + 2-oxoglutarate + O2
?
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r
100-amino acid alpha1(I) collagen fragment L-proline + 2-oxoglutarate + O2
100-amino acid alpha1(I) collagen fragment trans-4-hydroxy-L-proline + succinate + CO2
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?
2-oxoglutarate + O2 + ascorbate
succinate + dehydroascorbate + CO2 + H2O
510-amino acid alpha1(I) collagen fragment + 2-oxoglutarate + O2
?
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r
510-amino acid alpha1(I) collagen fragment L-proline + 2-oxoglutarate + O2
510-amino acid alpha1(I) collagen fragment trans-4-hydroxy-L-proline + succinate + CO2
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?
Arg-Gly-(Pro-Pro-Gly)5 + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
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?
Argonaute 2 + 2-oxoglutarate + O2
4-hydroxyproline-Argonaute 2 + succinate + CO2
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regulation of Ago 2 protein activity via substrate protein stability involving Ago Pro700 residue, the Ago protein mutant P700A is destabilized, overview
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?
Argonaute protein + 2-oxoglutarate + O2
4-hydroxyproline-Argonaute protein + succinate + CO2
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i.e. GERp95 or Golgi endoplasmic reticulum protein 95 kDa. Recombinant substrate proteins expressed in HeLa S3 cells. Ago2 is a cytoplasmically exposed, peripheral membrane protein that exists in a protease-resistant complex. Hydroxylation of Pro700, which is important for Ago2 stability, but not of Ago1 or Ago3 stability, identification by mass spectrometric analysis. In vitro, both Ago2 and Ago4 seem to be more efficiently hydroxylated than Ago1 and Ago3 by recombinant human isozyme C-P4H(I)
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?
Asp-Ala-Leu-Thr-Leu-Leu-Ala-Pro-Ala-Ala-Gly-Asp-Thr-Ile-Ile-Ser-Leu-Phe-Gly + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
peptide representing hypoxia-inducible transcription factor alpha sequences
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?
Asp-Leu-Asp-Leu-Glu-Met-Leu-Ala-Pro-Tyr-Ile-Pro-Met-Asp-Asp-Asp-Phe-Gln-Leu + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
peptide representing hypoxia-inducible transcription factor alpha sequences
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?
biotin-Ahx-DLDLEALAP564YIPADDDFQL + 2-oxoglutarate + O2
?
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19mer HIF peptide not containing cysteine is used, in order to rule out an involvement of the cysteine nitrosylation
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?
bradykinin + 2-oxoglutarate + O2
4-hydroxyproline containing bradykinin
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?
carboxymethylated collagen L-proline + 2-oxoglutarate + O2
carboxymethylated collagen-trans-4-hydroxy-L-proline + succinate + CO2
collagen + 2-oxoglutarate + O2
4-hydroxyproline containing collagen + succinate + CO2
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?
DALTLLAPAAGDTIISLDYG + 2-oxoglutarate + O2
?
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NODD peptide derived from native HIF-1alpha395-414
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?
DALTLLAPAAGDTIISLFG + 2-oxoglutarate + O2
DALTLLA-((4R)-4-hydroxy-L-proline)-AAGDTIISLFG + succinate + CO2
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?
dansyl-Gly-Phe-Pro-Gly-OEt + 2-oxoglutarate + O2
dansyl-Gly-Phe-4-hydroxy-L-Pro-Gly-OEt + succinate + CO2
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?
dansyl-Gly-Phe-Pro-Gly-OEt + 2-oxoglutarate + O2
dansyl-Gly-Phe-trans-4-hydroxy-L-proline-Gly-OEt + succinate + CO2
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?
DLDLEALAPYIPADDDFQL + 2-oxoglutarate + O2
?
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CODD peptide derived from native HIF-1alpha556-574. PHD2 preferred CODD by 20fold over NODD
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?
DLDLEMLAPAIPMDDDFQL + 2-oxoglutarate + O2
DLDLEMLA-((4R)-4-hydroxy-L-proline)-AIPMDDDFQL + succinate + CO2
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?
DLDLEMLAPAIPMDDDFQLRSFDQ + 2-oxoglutarate + O2
DLDLEMLAPAIPMDDDFQLRSFDQ trans-4-hydroxy-L-proline + succinate + CO2
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?
DLDLEMLAPGIPMDDDFQL + 2-oxoglutarate + O2
DLDLEMLAPGIPMDDDFQL trans-4-hydroxy-L-proline + succinate + CO2
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?
DLDLEMLAPYIPMD + 2-oxoglutarate + O2
DLDLEMLAPYIPMD trans-4-hydroxy-L-proline + succinate + CO2
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?
DLDLEMLAPYIPMDD + 2-oxoglutarate + O2
DLDLEMLAPYIPMDD trans-4-hydroxy-L-proline + succinate + CO2
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?
DLDLEMLAPYIPMDDDF + 2-oxoglutarate + O2
DLDLEMLAPYIPMDDDF trans-4-hydroxy-L-proline + succinate + CO2
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?
DLDLEMLAPYIPMDDDFQL + 2-oxoglutarate + O2
DLDLEMLAPYIPMDDDFQL trans-4-hydroxy-L-proline + succinate + CO2
-
-
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?
DLDLEMLAPYIPMDDDFQLRSFDQ + 2-oxoglutarate + O2
DLDLEMLAPYIPMDDDFQLRSFDQ trans-4-hydroxy-L-proline + succinate + CO2
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-
?
DLEMLAPYIPMDDDFQL + 2-oxoglutarate + O2
DLEMLA-((4R)-4-hydroxy-L-proline)-YIPMDDDFQL + succinate + CO2
-
-
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?
EMLAPYIPMDDDFQL + 2-oxoglutarate + O2
EMLAPYIPMDDDFQL trans-4-hydroxy-L-proline + succinate + CO2
-
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?
Glu-Gly-(Pro-Pro-Gly)5 + 2-oxoglutarate + O2
4-hydroxyproline containing peptide + succinate + CO2
-
-
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?
Gly-Pro8 + 2-oxoglutarate + O2
4-hydroxyproline containing peptide
Gly-Val-Pro-Gly-Val + 2-oxoglutarate + O2
Gly-Val-4-hydroxyproline-Gly-Val + succinate + CO2
-
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?
human procollagen-like (P-P-G)10 peptide L-proline + 2-oxoglutarate + O2
human procollagen-like (P-P-G)10 peptide trans-4-hydroxy-L-proline + succinate + CO2
Q81LZ8
4 different peptide sequences, anaerobic conditions
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-
?
human procollagen-like (P-P-G)5 peptide L-proline + 2-oxoglutarate + O2
human procollagen-like (P-P-G)5 peptide trans-4-hydroxy-L-proline + succinate + CO2
Q81LZ8
5 different peptide sequences, anaerobic conditions
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-
?
hypoxia-induced factor alpha oxygen-dependent degradation domain L-proline + 2-oxoglutarate + O2
hypoxia-induced factor alpha oxygen-dependent degradation domain L-4-hydroxyproline + succinate + CO2
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in vitro hydroxylation of two proline residues, with preference for C-terminal proline
-
?
hypoxia-inducible factor alpha L-proline + 2-oxoglutarate + O2
hypoxia-inducible factor alpha trans-4-hydroxy-L-proline + succinate + CO2
hypoxia-inducible factor L-proline + 2-oxoglutarate + O2
hypoxia inducible factor trans-4-hydroxy-L-proline + succinate + CO2
hypoxia-inducible factor L-proline + 2-oxoglutarate + O2
hypoxia-inducible factor trans-4-hydroxy-L-proline + succinate + CO2
-
-
-
-
?
hypoxia-inducible transcription factor + 2-oxoglutarate + O2
hypoxia-inducible transcription factor trans-4-hydroxy-L-proline + succinate + CO2
L-Ala-(L-Pro)4-Gly-L-Ile-L-Pro-Gly-L-Tyr-(L-Pro)2-L-Ala-(L-Pro)4-Gly-L-Val-Gly-(L-Pro)4-L-Gln-Gly L-proline + 2-oxoglutarate + O2
L-Ala-(L-Pro)4-Gly-L-Ile-L-Pro-Gly-L-Tyr-(L-Pro)2-L-Ala-(L-Pro)4-Gly-L-Val-Gly-(L-Pro)4-L-Gln-Gly trans-4-hydroxy-L-proline + succinate + CO2
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-
peptide substrate representing proline-rich proteins expressed in the pharyngeal gland cells. 380% of the activity with (L-Pro-L-Glu-L-Pro-L-Pro-L-Ala)5 L-proline
-
?
L-Pro-L-Ala-L-Pro-L-Lys + 2-oxoglutarate + O2
L-Pro-L-Ala-L-Pro-L-Lys-trans-4-hydroxy-L-proline + succinate + CO2
-
-
-
-
r
L-Pro-L-Ile-(L-Pro)7-L-Leu-L-Pro-L-Gln-L-Asn-L-Leu-L-Ser-Gly-L-Ala-L-Pro L-proline + 2-oxoglutarate + O2
L-Pro-L-Ile-(L-Pro)7-L-Leu-L-Pro-L-Gln-L-Asn-L-Leu-L-Ser-Gly-L-Ala-L-Pro trans-4-hydroxy-L-proline + succinate + CO2
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peptide substrate representing proline-rich proteins expressed in the pharyngeal gland cells. 610% of the activity with (L-Pro-L-Glu-L-Pro-L-Pro-L-Ala)5 L-proline
-
?
L-Pro-L-Pro-Gly + 2-oxoglutarate + O2
L-Pro-L-Pro-Gly-trans-4-hydroxy-L-proline + succinate + CO2
L-proline + 2-oxoglutarate + O2
4-hydroxyproline + succinate + CO2
LAPYIPMDDDFQL + 2-oxoglutarate + O2
LAPYIPMDDDFQL trans-4-hydroxy-L-proline + succinate + CO2
-
-
-
?
Lys-Pro-Ala + 2-oxoglutarate + O2
Lys-4-hydroxyproline-Ala + succinate + CO2
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recombinant enzyme
-
?
lysine hydroxylated protocollagen + 2-oxoglutarate + O2
lysine 4-hydroxyproline containing protocollagen + succinate + CO2
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?
octa-L-proline + 2-oxoglutarate + O2
4-hydroxyproline containing peptide
PEG-Gly-Tyr-4-fluoroproline-GlyOEt + 2-oxoglutarate + O2
?
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-
-
-
?
peptidyl L-proline + 2-oxoglutarate + O2
peptidyl-trans-4-hydroxy-L-proline + succinate + CO2
-
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-
-
r
poly L-proline + 2-oxoglutarate + O2
poly trans-4-hydroxy-L-proline + succinate + CO2
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molecular weights of 5000 Da, 8000 Da and 32000 Da, respectively
-
-
?
poly(ethylene glycol)-Gly-L-Tyr-(2S)-4-thiaproline-Gly-OCH2CH2OH + 2-oxoglutarate + O2
poly(ethylene glycol)-Gly-L-Tyr-L-(2S,4R)-thiaoxoproline-Gly-OCH2CH2OH + ? + CO2
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6% of the activity with L-prolyl peptide substrate
-
-
?
poly(ethylene glycol)-Gly-L-Tyr-L-(2S,4S)-4-fluoroproline-Gly-OCH2CH2OH + 2-oxoglutarate + O2
poly(ethylene glycol)-Gly-L-Tyr-L-(2S)-4-oxoproline-Gly-OCH2CH2OH + ? + CO2
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26% of the activity with L-prolyl peptide substrate
-
-
?
poly(ethylene glycol)-Gly-L-Tyr-L-Pro-Gly-OCH2CH2OH + 2-oxoglutarate + O2
poly(ethylene glycol)-Gly-L-Tyr-L-(2S,4R)-4-hydroxy-Pro-Gly-OCH2CH2OH + succinate + CO2
-
-
-
-
?
poly(L-Pro) + 2-oxoglutarate + O2
poly(4-hydroxyproline) + succinate + CO2