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Information on EC 1.4.3.13 - protein-lysine 6-oxidase
for references in articles please use BRENDA:EC1.4.3.13
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EC Tree 1 Oxidoreductases
1.4 Acting on the CH-NH2 group of donors
1.4.3 With oxygen as acceptor
1.4.3.13 protein-lysine 6-oxidase
IUBMB Comments
Also acts on protein 5-hydroxylysine. This enzyme catalyses the final known enzymic step required for collagen and elastin cross-linking in the biosynthesis of normal mature extracellular matrices . These reactions play an important role for the development, elasticity and extensibility of connective tissue. The enzyme is also active on free amines, such as cadaverine or benzylamine [4,5]. Some isoforms can also use [protein]-N(6)-acetyl-L-lysine as substrate deacetamidating the substrate .
The expected taxonomic range for this enzyme is: Eukaryota, Archaea
- 1.4.3.13
- collagen
- fibrosis
- metastasis
- fiber
- copper
-
crosslinking
- amine
-
elastic
- glaucoma
- aortic
-
stiff
- beta-aminopropionitrile
- tropoelastin
-
copper-dependent
-
fibril
- metalloproteinases
- procollagens
-
biomechanical
- propeptide
-
exfoliation
-
fibrillar
-
tensile
- myofibroblasts
- aneurysm
-
pseudoexfoliation
- cutis
-
copper-binding
- fibrillin-1
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open-angle
- menkes
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copper-deficient
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prolapse
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fibrillogenesis
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cross-linkage
- desmosine
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copper-containing
- medicine
- benzylamine
- diagnostics
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pentosidine
- synthesis
-
microfibrillar
-
microfibrils
-
elastosis
- ehlers-danlos
-
elastolytic
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col3a1
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semicarbazide-sensitive
-
cuproenzymes
-
c-proteinase
-
fibulins
- hydroxylysine
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pyridinoline
Reaction Schemes
showSynonyms
lysyl oxidase, loxl2, loxl1, loxl4, loxl3, lox-pp, lysyl oxidases, lysyl oxidase-like 1, lysyl oxidase-like 2, lysyl oxidase-like, more
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SYNONYM 
ORGANISM
UNIPROT
COMMENTARY 
LITERATURE
LO
LOX
LOXL
LOXL1
Loxl2
LOXL3-sv1
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LOXL3 encodes two variants, LOXL3 and LOXL3-sv1, both of which function as amine oxidases with distinct tissue and substrate specificities from one another
lysyl oxidase
lysyl oxidase propetide
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has biological activity as a tumor suppressor, and it inhibits Erk1/2 Map kinase activation
lysyl oxidase-like 1
lysyl oxidase-like 2
lysyl oxidase-like protein
RAS excision protein
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-
-
-
additional information
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enzymes belong to the lysyl oxidase family of copper amine oxidases
LOX
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-
LOX
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LOX
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LOXactivity is essential to generate optimal chemotactic sensitivity of cells to chemoattractants by oxidizing specific cell surface proteins, such as platelet-derived growth factor-beta
LOX
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LOXactivity is essential to generate optimal chemotactic sensitivity of cells to chemoattractants by oxidizing specific cell surface proteins, such as platelet-derived growth factor-beta
LOXL1
-
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lysyl oxidase
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amine oxidase, oxidative deamination
lysyl oxidase
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lysyl oxidase
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-
lysyl oxidase-like 1
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REACTION
REACTION DIAGRAM
COMMENTARY
ORGANISM
UNIPROT
LITERATURE
[protein]-L-lysine + O2 + H2O = [protein]-(S)-2-amino-6-oxohexanoate + NH3 + H2O2
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-
-
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[protein]-L-lysine + O2 + H2O = [protein]-(S)-2-amino-6-oxohexanoate + NH3 + H2O2
information about the activation mechanism of copper
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[protein]-L-lysine + O2 + H2O = [protein]-(S)-2-amino-6-oxohexanoate + NH3 + H2O2
preprotein, lysyl oxidase gene family, amine oxidation through ping pong bi ter kinetic mechanism
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[protein]-L-lysine + O2 + H2O = [protein]-(S)-2-amino-6-oxohexanoate + NH3 + H2O2
Also acts on protein 5-hydroxylysine
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-
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[protein]-L-lysine + O2 + H2O = [protein]-(S)-2-amino-6-oxohexanoate + NH3 + H2O2
active site structure, a unique funnel defines substrate specificity
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[protein]-L-lysine + O2 + H2O = [protein]-(S)-2-amino-6-oxohexanoate + NH3 + H2O2
ping-pong mechanism of 2 half-reactions completing the catalytic cycle, overview
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[protein]-L-lysine + O2 + H2O = [protein]-(S)-2-amino-6-oxohexanoate + NH3 + H2O2
ping-pong mechanism of 2 half-reactions completing the catalytic cycle, overview
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[protein]-L-lysine + O2 + H2O = [protein]-(S)-2-amino-6-oxohexanoate + NH3 + H2O2
ping-pong mechanism of 2 half-reactions completing the catalytic cycle, overview
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REACTION TYPE
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
oxidation
-
-
-
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oxidative deamination
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-
-
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redox reaction
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-
-
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reduction
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-
-
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SYSTEMATIC NAME
IUBMB Comments
protein-L-lysine:oxygen 6-oxidoreductase (deaminating)
Also acts on protein 5-hydroxylysine. This enzyme catalyses the final known enzymic step required for collagen and elastin cross-linking in the biosynthesis of normal mature extracellular matrices [4]. These reactions play an important role for the development, elasticity and extensibility of connective tissue. The enzyme is also active on free amines, such as cadaverine or benzylamine [4,5]. Some isoforms can also use [protein]-N(6)-acetyl-L-lysine as substrate deacetamidating the substrate [6].
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CAS REGISTRY NUMBER
COMMENTARY
99676-44-5
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SUBSTRATE
PRODUCT
REACTION DIAGRAM
ORGANISM
UNIPROT
LITERATURE
COMMENTARY
Reversibility
r=reversible
ir=irreversible
?=not specified
r=reversible
ir=irreversible
?=not specified
benzylamine + O2 + H2O
benzaldehyde + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
collagen + O2 + H2O
?
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Substrates: collagen type I, II, III, IV, VI, VIII and X
Products: -
Products: -
?
collagen 2a1 + O2 + H2O
?
Substrates: -
Products: -
Products: -
?
collagen III + O2 + H2O
allysyl-collagen III + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
dichlorodihydrofluorescein diacetate + O2 + H2O
? + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
histone H2 + O2 + H2O
?
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Substrates: recombinant mature LOX and C-terminally deleted enzyme
Products: -
Products: -
?
peptidyl-L-hydroxylysyl-peptide + O2 + H2O
peptidyl-L-2-amino-5-hydroxy-6-oxohexanoyl-peptide + NH3 + H2O2
tropoelastin + ?
?
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Substrates: LOXL1 catalyzes the polymerization of tropoelastin
Products: -
Products: -
?
tropoelastin + O2 + H2O
? + H2O2 + NH3
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Substrates: oxidation of L-lysine residues, substrate from mammalia
Products: -
Products: -
?
[elastin]-L-lysine + O2 + H2O
[elastin]-(S)-2-amino-6-oxohexanoate + NH3 + H2O2
Substrates: the enzyme catalyzes the oxidative deamination of lysine residues of extracellular matrix proteins such as elastins and collagens and generate aldehyde groups. The oxidative deamination of lysine represents the foundational step for the cross-linking of elastin and collagen and thus is crucial for modeling of extracellular matrix
Products: -
Products: -
?
collagen + H2O + O2
allysyl-collagen + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
collagen + H2O + O2
allysyl-collagen + NH3 + H2O2
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Substrates: the enzyme initiates the enzymatic stage of collagen and elastin cross-linking. Lysyl oxidase activity is essential for the integrity maintenance of the dermis and for the homeostasis of the epidermis. The LOX protein plays a role in the skin carcinomas and invasion but not through its enzymatic activity
Products: -
Products: -
?
elastin + H2O + O2
allysyl-elastin + NH3 + H2O2
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Substrates: the enzyme initiates the enzymatic stage of collagen and elastin cross-linking. Lysyl oxidase activity is essential for the integrity maintenance of the dermis and for the homeostasis of the epidermis. The LOX protein plays a role in the skin carcinomas and invasion but not through its enzymatic activity
Products: -
Products: -
?
histone H1 + O2 + H2O
?
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Substrates: L-lysine residues in histone H1
Products: -
Products: -
?
histone H1 + O2 + H2O
?
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Substrates: L-lysine residues in histone H1
Products: -
Products: -
?
histone H1 + O2 + H2O
?
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Substrates: recombinant mature LOX and C-terminally deleted enzyme
Products: -
Products: -
?
histone H1 + O2 + H2O
?
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Substrates: L-lysine residues in histone H1
Products: -
Products: -
?
lysine:tyrosine (4:1) heteropolymer + O2 + H2O
? + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
lysine:tyrosine (4:1) heteropolymer + O2 + H2O
? + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
n-alkylamine + O2 + H2O
aldehyde + NH3 + H2O2
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Substrates: synthetic substrates
Products: -
Products: -
?
n-alkylamine + O2 + H2O
aldehyde + NH3 + H2O2
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Substrates: synthetic substrates
Products: -
Products: -
?
n-alkylamine + O2 + H2O
aldehyde + NH3 + H2O2
-
Substrates: synthetic substrates
Products: -
Products: -
?
peptidyl-L-hydroxylysyl-peptide + O2 + H2O
peptidyl-L-2-amino-5-hydroxy-6-oxohexanoyl-peptide + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
peptidyl-L-hydroxylysyl-peptide + O2 + H2O
peptidyl-L-2-amino-5-hydroxy-6-oxohexanoyl-peptide + NH3 + H2O2
-
Substrates: part of tropocollagen and tropoelastin
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: deamination of lysyl and hydroxylysyl residues is part of the pyridinoline biosynthetic pathway, pyridinoline is involved in formation of intracellular and extracellular collagen cross-links responsible for aging of connective tissue
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: enzyme plays a critical role in the formation and repair of extracellular matrix, formation of cross-links in fibrillar collagen and elastin, only in fibrillar aggregates
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: in a variety of basic globular proteins
Products: i.e. peptidyl-L-alpha-aminoadipic-delta-semialdehyde
Products: i.e. peptidyl-L-alpha-aminoadipic-delta-semialdehyde
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: enzyme plays a critical role in the formation and repair of extracellular matrix, formation of cross-links in fibrillar collagen and elastin, only in fibrillar aggregates
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: key enzyme in formation of lysine cross-links in side collagen and elastin molecules
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: substrates are proteins of the extracellular matrix
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: in a variatey of basic globular proteins
Products: i.e. peptidyl-L-alpha-aminoadipic-delta-semialdehyde
Products: i.e. peptidyl-L-alpha-aminoadipic-delta-semialdehyde
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: in collagen and elastin fibrills
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: formation of cross-links in mature fibrillar collagen and elastin fibers in the extracellular space
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: initiation of covalent intermolecular cross-linking of collagen by the enzyme action on telopeptidyl lysine and hydroxylysine residues
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: lysyl oxidase plays a critical role in the formation of the extracellular matrix, and its activity is required for the normal maturation and cross-linking of collagen and elastin
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: enzyme plays a critical role in the formation and repair of extracellular matrix, formation of cross-links in fibrillar collagen and elastin, only in fibrillar aggregates
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: in a variatey of basic globular proteins
Products: i.e. peptidyl-L-alpha-aminoadipic-delta-semialdehyde
Products: i.e. peptidyl-L-alpha-aminoadipic-delta-semialdehyde
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-allysyl-peptide + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-L-allysyl-peptide + NH3 + H2O2
-
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-L-allysyl-peptide + NH3 + H2O2
-
Substrates: part of tropocollagen and tropoelastin
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-L-allysyl-peptide + NH3 + H2O2
-
Substrates: part of elastin and collagen
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-L-allysyl-peptide + NH3 + H2O2
Substrates: -
Products: -
Products: -
?
peptidyl-L-lysyl-peptide + O2 + H2O
peptidyl-L-allysyl-peptide + NH3 + H2O2
-
Substrates: crosslinkage in fibrillar collagen and elastin
Products: -
Products: -
?
[collagen]-L-lysine + O2 + H2O
[collagen]-(S)-2-amino-6-oxohexanoate + NH3 + H2O2
Substrates: -
Products: -
Products: -
?
[collagen]-L-lysine + O2 + H2O
[collagen]-(S)-2-amino-6-oxohexanoate + NH3 + H2O2
Substrates: the enzyme catalyzes the oxidative deamination of lysine residues of extracellular matrix proteins such as elastins and collagens and generate aldehyde groups. The oxidative deamination of lysine represents the foundational step for the cross-linking of elastin and collagen and thus is crucial for modeling of extracellular matrix
Products: -
Products: -
?
additional information
?
-
-
Substrates: enzyme modifies cellular activity and can function as a messenger shuttling information between the intra- and extracellular compartments
Products: -
Products: -
?
additional information
?
-
-
Substrates: enzyme contains a cytokine receptor domain, substrate specificity, preference for the L-lysine residues in the sequence N-acetyl-(Gly)4-Glu-Lys-(Gly)5-amide, enzyme is tightly associated to the substrates in connective tissue
Products: -
Products: -
?
additional information
?
-
-
Substrates: the enzyme does not affect smooth muscle cell proliferation nor elastin precursor, tropoelastin, synthesis, nor total elastin synthesis in a per-cell-basis
Products: -
Products: -
?
additional information
?
-
-
Substrates: critical role for lysyl oxidase in zebrafish notochord morphogenesis
Products: -
Products: -
?
additional information
?
-
Substrates: LOX induced chromatin changes, mechanism via histone deamination
Products: -
Products: -
?
additional information
?
-
Substrates: LOX induced chromatin changes, mechanism via histone deamination
Products: -
Products: -
?
additional information
?
-
Substrates: lysyl oxidase gene family. Enzyme seems to be involved in binding and crosslinking to other cell-surface and extracellular-matrix proteins
Products: -
Products: -
?
additional information
?
-
-
Substrates: lysyl oxidase gene family. Enzyme seems to be involved in binding and crosslinking to other cell-surface and extracellular-matrix proteins
Products: -
Products: -
?
additional information
?
-
-
Substrates: enzyme modifies cellular activity and can function as a messenger shuttling information between the intra- and extracellular compartments, enzyme expression in downregulated in Menkes disease, a disorder of copper transport
Products: -
Products: -
?
additional information
?
-
-
Substrates: LOX induced chromatin changes, mechanism via histone deamination
Products: -
Products: -
?
additional information
?
-
-
Substrates: the enzyme forms are differently involved in tumor suppression, cellular senescence, and chemotaxis
Products: -
Products: -
?
additional information
?
-
-
Substrates: enzyme contains a cytokine receptor domain, substrate specificity, preference for the L-lysine residues in the sequence N-acetyl-(Gly)4-Glu-Lys-(Gly)5-amide, enzyme is tightly associated to the substrates in connective tissue
Products: -
Products: -
?
additional information
?
-
-
Substrates: lysyl oxidase and 4 LOX-like proteins LOXL, LOXL2, LOXL3, and LOXL4 all contain a cytokine receptor-like domain
Products: -
Products: -
?
additional information
?
-
-
Substrates: lysyl oxidase and 4 LOX-like proteins LOXL, LOXL2, LOXL3, and LOXL4 all contain a cytokine receptor-like domain
Products: -
Products: -
?
additional information
?
-
-
Substrates: potential role of lysyl oxidase/placental lactogen interactions in breast epithelial cell proliferation and migration
Products: -
Products: -
?
additional information
?
-
-
Substrates: LOXL3 encodes two variants, LOXL3 and LOXL3-sv1, both of which function as amine oxidases with distinct tissue and substrate specificities
Products: -
Products: -
?
additional information
?
-
-
Substrates: catalyzes the cross-linking of collagens or elastin in the extracellular compartment, thereby regulating the tensile strength of tissues
Products: -
Products: -
?
additional information
?
-
-
Substrates: purifiedLOXreadily oxidizes peptidyl lysines in basic globular proteins with isoelectric points greater than pH 8, whereas no oxidation is detected in neutral or acidic proteins with isoelectric points less than pH 8
Products: -
Products: -
?
additional informatio