Information on EC 2.4.2.10 - orotate phosphoribosyltransferase

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The expected taxonomic range for this enzyme is: Bacteria, Eukaryota

EC NUMBER
COMMENTARY hide
2.4.2.10
-
RECOMMENDED NAME
GeneOntology No.
orotate phosphoribosyltransferase
REACTION
REACTION DIAGRAM
COMMENTARY hide
ORGANISM
UNIPROT
LITERATURE
orotidine 5'-phosphate + diphosphate = orotate + 5-phospho-alpha-D-ribose 1-diphosphate
show the reaction diagram
REACTION TYPE
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
pentosyl group transfer
-
-
-
-
PATHWAY
BRENDA Link
KEGG Link
MetaCyc Link
Drug metabolism - other enzymes
-
-
Metabolic pathways
-
-
pyrimidine metabolism
-
-
Pyrimidine metabolism
-
-
UMP biosynthesis
-
-
SYSTEMATIC NAME
IUBMB Comments
orotidine-5'-phosphate:diphosphate phospho-alpha-D-ribosyl-transferase
The enzyme from higher eukaryotes also catalyses the reaction listed as EC 4.1.1.23, orotidine-5'-phosphate decarboxylase.
CAS REGISTRY NUMBER
COMMENTARY hide
9030-25-5
-
ORGANISM
COMMENTARY hide
LITERATURE
UNIPROT
SEQUENCE DB
SOURCE
strain ATCC 6872
-
-
Manually annotated by BRENDA team
strain 1S-4
-
-
Manually annotated by BRENDA team
strain 1S-4
-
-
Manually annotated by BRENDA team
FCB strain
-
-
Manually annotated by BRENDA team
strain ATCC 17536. When grown on glucose as carbon source, supplementation with orotic acid significantly decreases orotate phosphoribosyltransferase and orotidine 5'-monophoshate decarboxylase activities. In a strain deficient for dihydroorotase activity, orotic acid induces both phosphoribosyltransferase and orotidine 5'-monophoshate decarboxylase activites
-
-
Manually annotated by BRENDA team
biovar trifolium
-
-
Manually annotated by BRENDA team
-
-
-
Manually annotated by BRENDA team
black gram
-
-
Manually annotated by BRENDA team
GENERAL INFORMATION
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
malfunction
-
decreased activity of OPRT plays an important role in the acquired resistance of gastric cancer cells towards 5-fluorouracil. OPRT-knockout MKN-45 parent cells using small interfering RNA demonstrate 15.8fold increased resistance to 5-fluorouracil compared to the control cell
SUBSTRATE
PRODUCT                       
REACTION DIAGRAM
ORGANISM
UNIPROT
COMMENTARY
(Substrate) hide
LITERATURE
(Substrate)
COMMENTARY
(Product) hide
LITERATURE
(Product)
Reversibility
r=reversible
ir=irreversible
?=not specified
5-fluoroorotate + 5-phospho-alpha-D-ribose 1-diphosphate
?
show the reaction diagram
-
-
-
?
5-fluoroorotate + phosphoribose diphosphate
5-fluoroorotidine 5'-phosphate + diphosphate
show the reaction diagram
-
-
-
?
5-fluorouracil + 5-phospho-alpha-D-ribose 1-diphosphate
5-fluorouridine 5'-phosphate + diphosphate
show the reaction diagram
orotate + 5-alpha-D-phosphorylribose 1-diphosphate
diphosphate + orotidine 5'-phosphate
show the reaction diagram
-
-
-
-
?
orotate + 5-phospho-alpha-D-ribose 1-diphosphate
orotidine 5'-phosphate + diphosphate
show the reaction diagram
orotate methylester + phosphoribose diphosphate
diphosphate + orotidine 4-methylester
show the reaction diagram
-
-
-
-
?
orotidine + diphosphate
orotate + alpha-D-ribose 1-diphosphate
show the reaction diagram
orotidine 5'-phosphate + diphosphate
orotate + 5-phospho-alpha-D-ribose 1-diphosphate
show the reaction diagram
orotidine 5'-phosphate + phosphonoacetic acid
?
show the reaction diagram
orotidine 5-phosphate + diphosphate
orotate + 5-phospho-alpha-D-ribose 1-diphosphate
show the reaction diagram
uracil + phosphoribose diphosphate
diphosphate + uridine
show the reaction diagram
-
-
-
-
?
additional information
?
-
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
5-fluorouracil + 5-phospho-alpha-D-ribose 1-diphosphate
5-fluorouridine 5'-phosphate + diphosphate
show the reaction diagram
orotate + 5-phospho-alpha-D-ribose 1-diphosphate
orotidine 5'-phosphate + diphosphate
show the reaction diagram
orotidine 5-phosphate + diphosphate
orotate + 5-phospho-alpha-D-ribose 1-diphosphate
show the reaction diagram
-
-
-
-
?
additional information
?
-
METALS and IONS
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
Ba2+
-
slight activation
INHIBITORS
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
1-deazaorotic acid
-
-
2,6-dihydroxypyridine-4-carboxylic acid
-
competitive to orotate, uncompetitive to 5-phospho-alpha-D-ribose 1-diphosphate
3-(2-hydroxybenzylidene)-2,6-dioxo-1,2,3,6-tetrahydropyridine-4-carboxylic acid
-
competitive to orotate, uncompetitive to 5-phospho-alpha-D-ribose 1-diphosphate
3-benzylidene-2,6-dioxo-1,2,3,6-tetrahydropyridine-4-carboxylic acid
-
competitive to orotate, uncompetitive to 5-phospho-alpha-D-ribose 1-diphosphate
5-aminoorotate
50% inhibition at 0.01 mM
5-azaorotic acid
-
-
5-Bromouracil
-
-
5-Chlorouracil
-
-
5-Fluoroorotate
5-fluorouracil
50% inhibition at 0.005-0.01 mM
5-Methylorotate
50% inhibition at 0.001 mM
5-phospho-alpha-D-ribose 1-diphosphate
6-hydroxy-5-((2-hydroxyethylamino)methyl)-2-oxo-1,2-dihydropyridine-4-carboxylic acid
-
competitive to orotate, uncompetitive to 5-phospho-alpha-D-ribose 1-diphosphate
adenine
adenosine
allopurinol
-
-
anthranilate
-
weak
arabinose 5-phosphate
-
at higher concentrations
Azauracil
-
-
Azauridine
-
-
Barbituric acid
-
-
Co2+
-
inhibits Mg2+-activation
cytosine
-
-
dihydroorotate
-
10 mM
diphosphate
dUMP
-
-
erythrose 4-phosphate
-
at higher concentrations
fructose 1-phosphate
-
at higher concentrations
fructose 6-phosphate
-
at higher concentrations
HgCl2
-
slight inhibition of mutant, not wild-type
iodoacetamide
-
slight inhibition of mutant, not wild-type
NEM
-
slight inhibition of mutant, not wild-type
nicotinate
-
weak
Orotate
Orotic acid
-
when grown on glucose as carbon source, supplementation with orotic acid significantly decreases orotate phosphoribosyltransferase and orotidine 5'-monophoshate decarboxylase activities. In a strain deficient for dihydroorotase activity, orotic acid induces both phosphoribosyltransferase and orotidine 5'-monophoshate decarboxylase activites
Orotidine
-
10 mM
orotidine 5'-phosphate
orotidylate
Oxipurinol
-
-
p-nitrophenyl beta-D-ribose
p-nitrophenyl beta-D-ribose 5'-phosphate
phosphate
pyrazofurin
50% inhibition at 0.006-0.024 mM
ribose 5-phosphate
sulfate
Uracil
uridine
-
-
xanthosine-5'-phosphate
-
-
additional information
ACTIVATING COMPOUND
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
dithiothreitol
-
stimulating
KM VALUE [mM]
SUBSTRATE
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
0.025
5-Fluoroorotate
pH 8.0, 37C
0.0016 - 10.01
5-phospho-alpha-D-ribose 1-diphosphate
0.013 - 0.25
diphosphate
0.002 - 0.44
Orotate
0.091 - 0.096
Orotidine
0.03
orotidine 5'-monophosphate
-
-
0.0015 - 0.045
orotidine 5'-phosphate
1.59 - 3.84
Phosphonoacetic acid
additional information
additional information
TURNOVER NUMBER [1/s]
SUBSTRATE
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
4
5-Fluoroorotate
Plasmodium falciparum
Q8N0R1
pH 8.0, 37C
3.2
5-phospho-alpha-D-ribose 1-diphosphate
Plasmodium falciparum
Q8N0R1
pH 8.0, 37C
0.28 - 1.3
diphosphate
0.6 - 10.8
Orotate
0.024 - 0.042
Orotidine
0.014 - 2.6
orotidine 5'-phosphate
0.017 - 0.023
Phosphonoacetic acid
kcat/KM VALUE [1/mMs-1]
SUBSTRATE
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
6 - 6.6
5-phospho-alpha-D-ribose 1-diphosphate
10
diphosphate
Mycobacterium tuberculosis
P9WHK9
ping-pong mechanism, pH 8.0, 25C
17
60 - 64
Orotate
0.26 - 0.44
Orotidine
4.8 - 530
orotidine 5'-phosphate
Ki VALUE [mM]
INHIBITOR
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
IMAGE
0.00047
1-deazaorotic acid
-
-
0.00025 - 0.0082
2,6-dihydroxypyridine-4-carboxylic acid
0.0074 - 0.014
3-(2-hydroxybenzylidene)-2,6-dioxo-1,2,3,6-tetrahydropyridine-4-carboxylic acid
0.00019 - 0.0065
3-benzylidene-2,6-dioxo-1,2,3,6-tetrahydropyridine-4-carboxylic acid
0.0021
5-azaorotic acid
-
-
0.0482
5-bromoorotate
pH 8.0, 37C
0.0954
5-Fluoroorotate
pH 8.0, 37C
0.0561
5-fluorouracil
pH 8.0, 37C
0.0153
5-iodoorotate
pH 8.0, 37C
0.0214
5-Methylorotate
pH 8.0, 37C
0.042 - 0.1501
5-phospho-alpha-D-ribose 1-diphosphate
0.0062 - 0.0135
6-hydroxy-5-((2-hydroxyethylamino)methyl)-2-oxo-1,2-dihydropyridine-4-carboxylic acid
0.026 - 0.209
diphosphate
0.0075 - 0.063
Orotate
0.00044 - 0.035
orotidine 5'-phosphate
0.0063 - 0.01
orotydilate
0.000121 - 0.000188
p-nitrophenyl beta-D-ribose
0.00004 - 0.000041
p-nitrophenyl beta-D-ribose 5'-phosphate
0.0705
pyrazofurin
pH 8.0, 37C
2.7 - 17.6
sulfate
0.1212
Uracil
pH 8.0, 37C
SPECIFIC ACTIVITY [µmol/min/mg]
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
0.0000045
-
untransfected TMK-1 cells
0.000009
-
untransfected MKN-45 cells
0.000072
-
transfected MKN-45-OPRT cells
0.000172
-
transfected TMK-OPRT cells
0.115
-
-
0.41
-
mutant
0.69
-
-
3 - 8
wild type enzyme
4.32
-
37 C, pH 8.0
15.2
-
wild-type
24
-
recombinant enzyme from crude cell extract, in the presence of Mg2+
45
-
in the absence of added OMP-decarboxylase, EC 4.1.1.23
65
-
in the presence of added OMP-decarboxylase, EC 4.1.1.23
71
-
recombinant enzyme after 17fold purification, in the presence of Co2+
225
-
recombinant enzyme after 17fold purification, in the presence of Mn2+
423
-
recombinant enzyme after 17fold purification, in the presence of Mg2+
additional information
pH OPTIMUM
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
5.5 - 6.5
-
reverse (pyrophosphorolysis) reaction
7 - 7.5
-
preferred buffer: Tris-HCl, when substrates are non-saturating, phosphate buffer, when substrate are saturating, not: maleate, imidazole, N-tris[hydroxymethyl] methyl-2-aminoethane-sulfonic acid (i.e. TES) or 3-[N-morpholino]-2-hydroxypropanesulfonic acid (i.e. MOPS) buffer
7.4
-
assay at
7.5
-
assay at
7.6
-
assay at
8.5
-
mutant
9.5
-
wild-type
10
-
recombinant enzyme
10.5 - 11.5
-
forward (phosphoribosyl transfer) reaction
additional information
-
2 isozymes: pI: 5.65 (minor form), pI: 5.85 (major form)
pH RANGE
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
6 - 8
recombinant monofunctional enzyme
7.2 - 9.5
-
about half-maximal activity at pH 7.2 and 9.5, mutant
7.2 - 9.2
-
about half-maximal activity at pH 7.2 and about 70% of maximal activity at pH 9.2
8.5 - 10.5
-
about half-maximal activity at pH 8.5 and about 70% of maximal activity at pH 10.5, wild-type
TEMPERATURE OPTIMUM
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
20
-
assay at
pI VALUE
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
5.4
-
predicted from amino acid sequence
SOURCE TISSUE
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
SOURCE
-
in normal non-neoplastic colorectal epithelium, granular staining is observed only in the crypt
Manually annotated by BRENDA team
-
in gallbladder carcinoma, mucosal neoplastic epithelium shows dense cytoplasmic expression but expression is absent in the deeply invasive lesions
Manually annotated by BRENDA team
-
mucosal neoplastic epithelium, granular expression
Manually annotated by BRENDA team
-
OPRT is involved in early events of invasion of hepatocellular carcinoma cells
Manually annotated by BRENDA team
-
normal tissue, hormone-sensitive prostate cancer (HSPC) or hormonerefractory prostate cancer (HRPC) tissue
Manually annotated by BRENDA team
-
gastric cancer cell line with low baseline expression levels of OPRT
Manually annotated by BRENDA team
additional information
LOCALIZATION
ORGANISM
UNIPROT
COMMENTARY hide
GeneOntology No.
LITERATURE
SOURCE
-
in metastatic lymph nodes and lymphovascularly invasive lesions
Manually annotated by BRENDA team
PDB
SCOP
CATH
ORGANISM
UNIPROT
Aeropyrum pernix (strain ATCC 700893 / DSM 11879 / JCM 9820 / NBRC 100138 / K1)
Burkholderia cenocepacia (strain ATCC BAA-245 / DSM 16553 / LMG 16656 / NCTC 13227 / J2315 / CF5610)
Corynebacterium diphtheriae (strain ATCC 700971 / NCTC 13129 / Biotype gravis)
Escherichia coli (strain K12)
Francisella tularensis subsp. tularensis (strain SCHU S4 / Schu 4)
Helicobacter pylori (strain ATCC 700392 / 26695)
Leishmania donovani (strain BPK282A1)
Saccharomyces cerevisiae (strain ATCC 204508 / S288c)
Saccharomyces cerevisiae (strain ATCC 204508 / S288c)
Saccharomyces cerevisiae (strain ATCC 204508 / S288c)
Saccharomyces cerevisiae (strain ATCC 204508 / S288c)
Saccharomyces cerevisiae (strain ATCC 204508 / S288c)
Salmonella typhimurium (strain LT2 / SGSC1412 / ATCC 700720)
Salmonella typhimurium (strain LT2 / SGSC1412 / ATCC 700720)
Salmonella typhimurium (strain LT2 / SGSC1412 / ATCC 700720)
Streptococcus mutans serotype c (strain ATCC 700610 / UA159)
Vibrio cholerae serotype O1 (strain ATCC 39315 / El Tor Inaba N16961)
MOLECULAR WEIGHT
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
39000
-
gel filtration
40000
-
gel filtration
45400
-
native enzyme, gel filtration
47000
-
K-12, gel filtration
50000
-
sucrose density gradient centrifugation
52000
-
Western blot analyis
140000
SUBUNITS
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
homodimer
monomer
-
1 * 51500, SDS-PAGE, enzyme activity and orotidine 5'-monophosphate decarboxylating activity on a single polypeptide
tetramer
additional information
Crystallization/COMMENTARY
ORGANISM
UNIPROT
LITERATURE
ligated with sulfate, homodimer
-
sitting-drop vapour-diffusion method, polyethylene glycol 300
-
sitting drop vapour diffusion method, the apoenzyme is crystallized in 100 mM Tris-HCl, pH 7.4, 0.2 M magnesium acetate, and 26% (w/v) PEG 6000, crystals of OPRTase in complex with substrates (5.0 mM 5-phospho-alpha-D-ribose 1-diphosphate and 5.0 mM orotate) or product (5.0 mM orotidine 5'-phosphate) are grown in 100 mM sodium acetate, pH 4.6, 0.14 M ammonium acetate, and 38% (w/v) PEG 4000, crystals of the OMP complex are grown from solutions containing 100 mM Tris HCl, pH 7.4, 0.08 M magnesium acetate, and 28% (w/v) PEG 6000
complexed with orotidine monophosphate, homodimer, with catalytically important residues from one subunit available for the other
-
hanging-grop vapour-diffusion method, 2.3 M ammonium sulfate, pH 8.6
pH STABILITY
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
4 - 9
-
stable
489712
7.5 - 9.5
-
at least 6 months stable, -20C
489715
TEMPERATURE STABILITY
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
50
-
OPRTase activity decreases with the increase in incubation temperature and more than 50% of the activity is lost upon incubation at 50C for 30 min
90
-
20 min, 70% of activity remaining
GENERAL STABILITY
ORGANISM
UNIPROT
LITERATURE
0.05 mM UMP and 1% polyethyleneglycol stabilize during final stage of purification
-
5 mM Mg2+, 1 mM phosphoribose diphosphate and 2 mM DTT stabilize dilute enzyme solutions
-
dialysis against phosphate buffer, pH 7 or 7.5, unstable to, 2 mM DTT protect
-
dilution inactivates
-
native enzyme complex, stable for at least 6 months at -20C or -80C in 50 mM Hepes, pH 7.4, 300 mM NaCl, 5 mM dithiothreitol, 20% glycerol. At 4C, loss of 50% of activity within 4 weeks.
phosphate, 0.3 M and above, stabilizes
-
proteins, e.g. albumin, do not stabilize dilute enzyme solutions
-
recombinant enzyme dimer, stable for at least 3 months at -20C or -80C in 50 mM Hepes, pH 7.4, 5 mM dithiothreitol, 20% glycerol. In absence of dithiothreitol and glycerol, loss of 50% of activity during overnight storage at 4C.
recombinant enzyme, stable for at least 3 months at 4C or -20C in presence of 1 mM dithiothreitol and 10% glycerol
STORAGE STABILITY
ORGANISM
UNIPROT
LITERATURE
-20C, concentrated enzyme solution, at least 4 months
-
-20C, pH 7.5-9.5, at least 6 months
-
-25C, in 50% v/v glycerol, 0.1 mM DTT, about 35% loss of activity within 40 days
-
-60C to-20C, crude enzyme or ammonium sulfate preparation, 2 years
-
-76C, at least 6 months
-
-80C, stable up to 12 months, 5% glycerol, pH 8.0
4C, 50 mM Tris-HCl, 2 mM EDTA, 2 mM mercaptoethanol, pH 6.4, several months, no apparent loss of activity
-
4C, gradual loss of activity at pH 7.5-9.5
-
4C, t1/2: 1.5 days
-
frozen, partially purified enzyme in phosphate buffer, pH 7 or 7.5, 2 months
-
Purification/COMMENTARY
ORGANISM
UNIPROT
LITERATURE
ammonium sulfate precipitation, DEAE-Sepharose Fast Flow column chromatography, and Superdex-75 gel filtration
-
anion exchange chromatography, gel filtration, hydrophobic interaction chromatography
from in vitro cultivation of organism
-
immobilized metal ion affinity chromatography (Co2+), His-tag cleavage (TEV protease), immobilized metal ion affinity chromatography (Co2+)
-
immobilized metal ion affinity chromatography (Ni2+)
-
immobilized metal ion affinity chromatography (Ni2+), gel filtration
Ni2+ affinity column
OMP-agarose affinity chromatography
-
partial, affinity chromatography on Blue-Dextran- or Cibacron Blue F3GA-Sepharose
-
partial, not separable from EC 4.1.1.23
-
partial, rapid tandem affinity column method
-
recombinant enzyme
Cloned/COMMENTARY
ORGANISM
UNIPROT
LITERATURE
expressed from pDEST14-OPRT in Escherichia coli BL21-AI competent cells
-
expressed in Escherichia coli
expressed in Escherichia coli BL21(DE3)
expressed in Escherichia coli DH5alpha cells
-
His-tagged protein expressed in Escherichia coli BL21(DE3)
His-tagged protein expressed in Escherichia coli BL21(DE3), gel filtration
-
into the pDONR221 vector, expressed from pDEST14-OPRT in Escherichia coli BL21-AI competent cells
-
mutants are expressed in Escherichia coli strain CS101-4UI
OPRT, stable overexpression in the gastric cancer cell lines TMK-1 and MKN-45, enhancing the effect of 5-fluorouracil on the cells, transfected cells show a similar growth curve as the wild-type parent cells, while the drug sensitivity is increased compared to the untransfected cells, overview
-
structural gene pyrE
-
structural gene pyrE, subcloned from a genomic library in pBR328(pAV002), transduced into pyr-auxotroph Salmonella typhimurium strain SA2434 with phage P22, finally cloned to and expressed in overproducing strain Escherichia coli MB13
-
EXPRESSION
ORGANISM
UNIPROT
LITERATURE
lexpression of OPRT in mucosal carcinoma components, in infiltrative components, in lymphovascularly invasive lesions and in metastatic lymph nodes. No significant correlation between OPRT expression and cancer stage, T-grade or N-grade, but OPRT expression correlates positively with the presence of lymphovascular invasion. OPRT is diffusely expressed to a greater extent, especially in the tumor cells adjacent to the infiltrative margin
-
low expression in normal prostate gland tissue
-
mRNA expression of OPRT is markedly decreased to 12.4% and activity to 46.9% in the 5-fluorouracil resistant MKN-45/F2R cell line compared to the MKN-45 parent cell line. OPRT gene expression is markedly decreased to 14.4% when down-regulated by siRNA
-
non-cancerous components do not express OPRT
-
strong expression of OPRT in prostate cancer cells, significant correlation between OPRT mRNA expression levels and the tumor pathological grade. The OPRT/dihydropyrimidine dehydrogenase expression ratio in the HRPC group is significantly higher than that in the low grade HSPC group
-
ENGINEERING
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
G213A
-
OPRT, the gene polymorphism predicts toxicity in patients treated with bolus 5-fluorouracil regimen, the Ala allele in the enzyme G213A polymorphism and the two tandem repeats in the TYMS promoter polymorphism are associated with grade 3 to 4 neutropenia and diarrhea, distribution in 69 patients samples, genotyping, relationship between OPRT mRNA expression and the OPRT G213A polymorphism, overview
D131A
severly reduced activity
D131A/D132A
severly reduced activity
D132A
severly reduced activity
D125N
-
active site of enzyme requires D125 of one subunit and K103 of second subunit
K103A
-
active site of enzyme requires D125 of one subunit and K103 of second subunit
additional information
APPLICATION
ORGANISM
UNIPROT
COMMENTARY hide
LITERATURE
diagnostics
medicine
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