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benzyloxycarbonyl-Arg p-nitrobenzyl ester + H2O
benzyloxycarbonyl-Arg + p-nitrobenzyl alcohol
-
-
-
?
benzyloxycarbonyl-L-Arg-thiobenzyl ester + H2O
benzyloxycarbonyl-L-Arg + thiobenzyl alcohol
-
-
-
-
?
benzyloxycarbonyl-L-Lys-thiobenzyl ester + H2O
benzyloxycarbonyl-L-Lys + thiobenzyl alcohol
-
-
-
?
benzyloxycarbonyl-Lys-thiobenzyl ester + H2O
benzyloxycarbonyl-Lys + thiobenzyl alcohol
-
-
-
-
?
beta-tubulin + H2O
?
cleaves beta-tubulin after Arg62 (YVPR-/-AV) and Arg282 (QQYR-/-AL)
-
-
?
Bid protein + H2O
tBid protein + ?
-
GzmK directly processes Bid to produce its active form tBid
-
-
?
Cbz-L-Arg-4-nitroanilide + H2O
Cbz-L-Arg + 4-nitroaniline
little hydrolyzed by Gr3
-
-
?
Cbz-L-Arg-7-amido-4-methylcoumarin + H2O
Cbz-L-Arg + 7-amino-4-methylcoumarin
little hydrolyzed by Gr3
-
-
?
Cbz-L-Lys-4-nitroanilide + H2O
Cbz-L-Lys + 4-nitroaniline
little hydrolyzed by Gr3
-
-
?
Cys-Gly-Tyr-Gly-Pro-Lys-Lys-Lys-Arg-Lys-Val-Gly-Gly + H2O
?
-
-
-
?
Cys-Gly-Tyr-Gly-Pro-Lys-Lys-Lys-Arg-Lys-Val-Gly-Gly + H2O
Cys-Gly-Tyr-Gly-Pro-Lys + Lys-Lys-Arg + Lys-Val-Gly-Gly
-
-
-
-
?
endoplasmic reticulum-associated SET complex + H2O
?
-
-
-
?
fluorescence resonance energy transfer substrate-25Arg peptide + H2O
?
most effectively hydrolyzes the substrate fluorescence resonance energy transfer substrate-25Arg peptide in the FRETS-25Xaa series tested
-
-
?
fluorescence resonance energy transfer substrate-25Lys peptide + H2O
?
far less effectively cleaved than fluorescence resonance energy transfer substrate-25Arg peptide
-
-
?
heterogeneous nuclear ribonucleoprotein K + H2O
?
granzyme K and granzyme A cleave with different kinetics at distinct sites
-
-
?
N-acetyl-YRFK-4-nitroanilide + H2O
N-acetyl-YRFK + 4-nitroaniline
-
-
-
-
?
N-tert-butoxycarbonyl-Ala-Ala-Asp-thiobenzyl ester
?
-
-
-
?
N-tert-butoxycarbonyl-Ala-Ala-Met-thiobenzyl ester
?
-
-
-
?
Nalpha-benzyloxycarbonyl-L-Arg-thiobenzyl ester
?
Nalpha-benzyloxycarbonyl-L-Lys-thiobenzyl ester
?
Nalpha-CBZ-L-lysine thiobenzyl ester
?
-
-
-
?
nucleosome assembly protein SET + H2O
?
-
-
-
-
?
redox factor-1/apurinic apyrimidinic endonuclease Ape1 + H2O
?
-
Ape1 cleavage by Granzyme K facilitates intracellular reactive oxygen species accumulation and enhances granzyme K-induced cell death
-
-
?
Z-Arg-thiobenzyl ester + H2O
Z-Arg + thiobenzyl alcohol
-
-
-
-
?
Z-Leu-Arg-Gly-Gly-7-amido-4-methylcoumarin + H2O
Z-Leu-Arg + Gly-Gly-7-amino-4-methylcoumarin
-
-
-
?
Z-Lys-thiobenzyl ester + H2O
?
-
-
-
?
Z-Lys-thiobenzyl ester + H2O
Z-Lys + thiobenzyl alcohol
-
-
-
-
?
additional information
?
-
Nalpha-benzyloxycarbonyl-L-Arg-thiobenzyl ester
?
-
-
-
?
Nalpha-benzyloxycarbonyl-L-Arg-thiobenzyl ester
?
-
-
-
?
Nalpha-benzyloxycarbonyl-L-Arg-thiobenzyl ester
?
-
-
-
?
Nalpha-benzyloxycarbonyl-L-Arg-thiobenzyl ester
?
-
-
-
?
Nalpha-benzyloxycarbonyl-L-Lys-thiobenzyl ester
?
-
-
-
?
Nalpha-benzyloxycarbonyl-L-Lys-thiobenzyl ester
?
-
-
-
?
Nalpha-benzyloxycarbonyl-L-Lys-thiobenzyl ester
?
-
-
-
?
Protein + H2O
?
-
-
-
?
additional information
?
-
-
cleaves synthetic thiobenzyl ester substrates after Lys and Arg, no substrates for the mature enzyme: Boc-Ala-Ala-Met-thiobenzyl ester and Boc-Ala-Ala-Asp-thiobenzyl ester
-
?
additional information
?
-
-
for single-residue thioester substrates the enzyme shows a 2fold preference for a Lys versus Arg residue at P1. With oligopeptide substrates the enzyme displays peptidolytic activity C-terminal to both Lys and Arg residues with comparable rates of hydrolysis
-
?
additional information
?
-
-
can trigger DNA fragmentation and is involved in apoptosis
-
?
additional information
?
-
most efficiently hydrolyzes the carboxylic side of Phe-Tyr-Arg (P3-P2-P1) sequence
-
-
?
additional information
?
-
-
most efficiently hydrolyzes the carboxylic side of Phe-Tyr-Arg (P3-P2-P1) sequence
-
-
?
additional information
?
-
granzyme K prefers Arg over Lys at P1 position. Granzyme K and granzyme A display highly restricted substrate specificities that overlapp only partially. Whereas granzyme K and granzyme A cleave SET with similar efficiencies likely at the same sites, both granzymes cleave the pre-mRNA-binding protein heterogeneous ribonuclear protein K with different kinetics at distinct sites. Granzyme K is markedly more efficient in cleaving heterogeneous ribonuclear protein K than granzyme A. Granzyme K, but not granzyme A, cleaves the microtubule network protein beta-tubulin after two distinct Arg residues. Neither GrK cleavage sites in beta-tubulin nor a peptide based proteomic screen reveal a clear granzyme K consensus sequence around the P1 residue, suggesting that GrK specificity depends on electrostatic interactions between exosites of the substrate and the enzyme
-
-
?
additional information
?
-
substrate identification analysing substrate repertoire and specificity by N-terminal COFRADIC assisted N-terminomics on the homologous enzyme. Comparisons of specificity profiles of human and mouse enzymes revealing only subtle differences and pointed to substrate occupancies in the P1, P1', and P2' position as the main determinants for substrate recognition, overview
-
-
?
additional information
?
-
-
substrate identification analysing substrate repertoire and specificity by N-terminal COFRADIC assisted N-terminomics on the homologous enzyme. Comparisons of specificity profiles of human and mouse enzymes revealing only subtle differences and pointed to substrate occupancies in the P1, P1', and P2' position as the main determinants for substrate recognition, overview
-
-
?
additional information
?
-
-
no substrates: Boc-Ala-Ala-Met-thiobenzyl ester and Boc-Ala-Ala-Asp-thiobenzyl ester
-
?
additional information
?
-
granzyme K expressing cytotoxic T lymphocytes protects against influenza virus in granzyme AB-/- mice. GrzK plays an important role in CD8+ T-cell cytotoxicity both in the presence and absence of grzA and B
-
-
?
additional information
?
-
-
substrate identification analysing substrate repertoire and specificity by N-terminal COFRADIC assisted N-terminomics on the homologous enzyme. Comparisons of specificity profiles of human and mouse enzymes revealing only subtle differences and pointed to substrate occupancies in the P1, P1', and P2' position as the main determinants for substrate recognition, overview
-
-
?
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3,3-diphenylpropanoyl-Pro-(4-AmPhGly)P(OPh)2
-
potent inhibitor
carbobenzoxy-Thr-(4-AmPhGly)P(OPh)2
-
quite specific but weak inhibitor
D-Phe-Pro-Arg-CH2Cl
-
good inhibitor
diisopropylfluorophosphate
-
complete inhibition at 0.1 mM
Glu-Gly-Arg-chloromethyl ketone
inter-alpha-inhibitor protein
physiological inhibitor, significantly inhibits catalytic activity in vitro
-
inter-alpha-trypsin inhibitor
-
complete inhibition at 66 nM
-
inter-alpha-trypsin inhibitor complex
-
physiologic inhibitor
-
Kunitz-type inhibitor aprotinin
-
-
-
L-Phe-L-Pro-L-Arg-chloromethyl ketone
-
complete inhibition at 0.1 mM
N-acetyl-YRFK-chloromethylketone
-
-
Phe-Pro-Arg-chloromethyl ketone
phenylmethylsulfonyl fluoride
Phenylmethylsulfonylfluoride
-
2 mM
second carboxy-terminal Kunitz-type domain of bikunin
-
Soybean trypsin inhibitor
-
50% inhibition at 0.025 mM
-
Tos-Lys-CH2Cl
-
complete inhibition at 0.1 mM
Z-LysP(OPh)2
-
best inhibitor
Z-Trp-Ch2Cl
-
moderate inhibitor
Zn2+
-
significant reduction of enzyme activity
3,4-dichloroisocoumarin
-
complete inactivation at 0.1 mM
3,4-dichloroisocoumarin
-
complete inhibition at 0.1 mM
4-aminobenzamidine
-
81% inactivation at 5 mM
4-aminobenzamidine
-
57% inactivation at 5 mM
Aprotinin
-
94% inactivation at 0.005 mM
Aprotinin
-
93% inactivation at 0.015 mM
Aprotinin
-
complete inhibition at 0.015 mM
Aprotinin
-
89% inactivation at 0.005 mM
benzamidine
-
50% inhibition at 27 mM
benzamidine
-
55% inactivation at 27 mM
benzamidine
-
56% inactivation at 27 mM
benzamidine
-
57% inactivation at 27 mM
bikunin
-
light chain of inter-alpha-trypsin inhibitor
-
bikunin
-
subunit of inter-alpha-trypsin inhibitor complex, physiologic inhibitor
-
bikunin
-
light chain of inter-alpha-trypsin inhibitor
-
Glu-Gly-Arg-chloromethyl ketone
-
97% inactivation at 0.1 mM
Glu-Gly-Arg-chloromethyl ketone
-
52% inactivation at 0.1 mM
leupeptin
-
50% inhibition at 0.2 mM
leupeptin
-
92% inactivation at 0.2 mM
leupeptin
-
22% inactivation at 0.2 mM
leupeptin
-
50% inhibition at 0.2 mM
leupeptin
-
22% inactivation at 0.2 mM
PefablocSC
-
44% inactivation at 1 mM
PefablocSC
-
94% inactivation at 1 mM
PefablocSC
-
94% inactivation at 1 mM
Phe-Pro-Arg-chloromethyl ketone
-
98% inactivation at 0.1 mM
Phe-Pro-Arg-chloromethyl ketone
-
complete inactivation at 0.1 mM
Phe-Pro-Arg-chloromethyl ketone
-
complete inactivation at 0.1 mM
phenylmethylsulfonyl fluoride
-
complete inactivation at 2 mM
phenylmethylsulfonyl fluoride
-
complete inhibition at 2 mM
second carboxy-terminal Kunitz-type domain of bikunin
-
-
-
second carboxy-terminal Kunitz-type domain of bikunin
-
physiologic inhibitor
-
second carboxy-terminal Kunitz-type domain of bikunin
-
-
-
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Aortic Aneurysm, Abdominal
Granzyme K - A novel marker to identify the presence and rupture of abdominal aortic aneurysm.
Arthritis
Inflammatory Gene Expression Profile and Defective Interferon-? and Granzyme K in Natural Killer Cells From Systemic Juvenile Idiopathic Arthritis Patients.
Arthritis, Juvenile
Inflammatory Gene Expression Profile and Defective Interferon-? and Granzyme K in Natural Killer Cells From Systemic Juvenile Idiopathic Arthritis Patients.
Asthma
Decrease of cytotoxic T cells in allergic asthma correlates with total serum immunglobulin E.
Colorectal Neoplasms
Clonally expanded EOMES+ Tr1-like cells in primary and metastatic tumors are associated with disease progression.
Hepatitis C
Engineering of TEV protease variants by yeast ER sequestration screening (YESS) of combinatorial libraries.
Hepatitis C
Yeast Endoplasmic Reticulum Sequestration Screening for the Engineering of Proteases from Libraries Expressed in Yeast.
Infections
Chlamydia trachomatis serovar L2 infection model using human lymphoid Jurkat cells.
Influenza, Human
Granzyme K expressing cytotoxic T lymphocytes protects against influenza virus in granzyme AB-/- mice.
Influenza, Human
Granzyme K inhibits replication of influenza virus through cleaving the nuclear transport complex importin ?1/? dimer of infected host cells.
Leukemia
Purification and cloning of a novel serine protease, RNK-Tryp-2, from the granules of a rat NK cell leukemia.
Lung Diseases
Granzyme K: a novel mediator in acute airway inflammation.
Lung Neoplasms
Clonally expanded EOMES+ Tr1-like cells in primary and metastatic tumors are associated with disease progression.
Lymphoma
A natural killer cell granule protein that induces DNA fragmentation and apoptosis.
Melanoma
Proteomic analysis and the antimetastatic effect of N-(4-methyl)phenyl-O-(4-methoxy) phenyl-thionocarbamate-induced apoptosis in human melanoma SK-MEL-28 cells.
Multiple Sclerosis
Unexpected role for granzyme K in CD56bright NK cell-mediated immunoregulation of multiple sclerosis.
Neoplasms
Anti-PD-1 monoclonal antibody MEDI0680 in a phase I study of patients with advanced solid malignancies.
Neoplasms
CD8-alpha T-cell infiltration in human papillomavirus-related oropharyngeal carcinoma correlates with improved patient prognosis.
Neoplasms
Human Asymptomatic Epitopes Identified from the Herpes Simplex Virus Tegument Protein VP13/14 (UL47) Preferentially Recall Polyfunctional Effector Memory CD44high CD62Llow CD8+ TEM Cells and Protect Humanized HLA-A*02:01 Transgenic Mice against Ocular Herpesvirus Infection.
Neoplasms
Ignition of p53 bomb sensitizes tumor cells to granzyme K-mediated cytolysis.
Neoplasms
Inflammatory Gene Expression Profile and Defective Interferon-? and Granzyme K in Natural Killer Cells From Systemic Juvenile Idiopathic Arthritis Patients.
Neoplasms
Valosin-containing protein cleavage by granzyme K accelerates an endoplasmic reticulum stress leading to caspase-independent cytotoxicity of target tumor cells.
Pneumonia
Granzyme K activates protease-activated receptor-1.
Sepsis
Granzyme K activates protease-activated receptor-1.
Typhoid Fever
Expression of intra- and extracellular granzymes in patients with typhoid fever.
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Babe, L.M.; Yoast, S.; Dreyer, M.; Schmidt, B.F.
Heterologous expression of human granzyme K in Bacillus subtilis and characterization of its hydrolytic activity in vitro
Biotechnol. Appl. Biochem.
27
117-124
1998
Homo sapiens
brenda
Wilharm, E.; Tschopp, J.; Jenne, D.E.
Biological activities of granzyme K are conserved in the mouse and account for residual Z-Lys-SBzl activity in granzyme A-deficient mice
FEBS Lett.
459
139-142
1999
Homo sapiens, Mus musculus
brenda
Wilharm, E.; Parry, M.A.; Friebel, R.; Tschesche, H.; Matschiner, G.; Sommerhoff, C.P.; Jenne, D.E.
Generation of catalytically active granzyme K from Escherichia coli inclusion bodies and identification of efficient granzyme K inhibitors in human plasma
J. Biol. Chem.
274
27331-27337
1999
Homo sapiens
brenda
Hink-Schauer, C.; Estebanez-Perpina, E.; Wilharm, E.; Fuentes-Prior, P.; Klinkert, W.; Bode, W.; Jenne, D.E.
The 2.2-A crystal structure of human pro-granzyme K reveals a rigid zymogen with unusual features
J. Biol. Chem.
277
50923-50933
2002
Homo sapiens
brenda
Shi, L.; Kam, C.M.; Powers, J.C.; Aebersold, R.; Greenberg, A.H.
Purification of three cytotoxic lymphocyte granule serine proteases that induce apoptosis through distinct substrate and target cell interactions
J. Exp. Med.
176
1521-1529
1992
Rattus norvegicus
brenda
Hameed, A.; Lowrey, D.M.; Lichtenheld, M.; Podack, E.R.
Characterization of three serine esterases isolated from human IL-2 activated killer cells
J. Immunol.
141
3142-3147
1988
Homo sapiens
brenda
Sayers, T.J.; Wiltrout, T.A.; Smyth, M.J.; Ottaway, K.S.; Pilaro, A.M.; Sowder, R.; Henderson, L.E.; Sprenger, H.; Lloyd, A.R.
Purification and cloning of a novel serine protease, RNK-Tryp-2, from the granules of a rat NK cell leukemia
J. Immunol.
152
2289-2297
1994
Rattus norvegicus
brenda
Jackson, D.S.; Fraser, S.A.; Ni, L.M.; Kam, C.M.; Winkler, U.; Johnson, D.A.; Froelich, C.J.; Hudig, D.; Powers, J.C.
Synthesis and evaluation of diphenyl phosphonate esters as inhibitors of the trypsin-like granzymes A and K and mast cell tryptase
J. Med. Chem.
41
2289-2301
1998
Homo sapiens
brenda
Bratke, K.; Kuepper, M.; Bade, B.; Virchow, J.C.; Luttmann, W.
Differential expression of human granzymes A, B, and K in natural killer cells and during CD8+ T cell differentiation in peripheral blood
Eur. J. Immunol.
35
2608-2616
2005
Homo sapiens
brenda
Bade, B.; Lohrmann, J.; ten Brinke, A.; Wolbink, A.M.; Wolbink, G.J.; ten Berge, I.J.; Virchow, J.C.; Luttmann, W.; Hack, C.E.
Detection of soluble human granzyme K in vitro and in vivo
Eur. J. Immunol.
35
2940-2948
2005
Homo sapiens
brenda
Babe, L.M.; Schmidt, B.F.
Granzyme K
Handbook of Proteolytic Enzymes (Barrett, A. J. , Rawlings, N. D. , Woessner; j. F. , eds. )Academic Press
2
1552-1554
2004
Homo sapiens, Rattus norvegicus
-
brenda
Babe, L.M.; Schmidt, B.F.
Granzyme K
Handbook of Proteolytic Enzymes (Barrett,A. J. ;Rawlings,N. D. ,Woessner)
2
1552-1554
2004
Mus musculus
-
brenda
Hirata, Y.; Inagaki, H.; Shimizu, T.; Li, Q.; Nagahara, N.; Minami, M.; Kawada, T.
Expression of enzymatically active human granzyme 3 in Escherichia coli for analysis of its substrate specificity
Arch. Biochem. Biophys.
446
35-43
2006
Homo sapiens (P49863), Homo sapiens
brenda
Zhao, T.; Zhang, H.; Guo, Y.; Zhang, Q.; Hua, G.; Lu, H.; Hou, Q.; Liu, H.; Fan, Z.
Granzyme K cleaves the nucleosome assembly protein SET to induce single-stranded DNA nicks of target cells
Cell Death Differ.
14
489-499
2007
Homo sapiens
brenda
Zhao, T.; Zhang, H.; Guo, Y.; Fan, Z.
Granzyme K directly processes Bid to release cytochrome c and endonuclease G leading to mitochondria-dependent cell death
J. Biol. Chem.
282
12104-12111
2007
Homo sapiens
brenda
Guo, Y.; Chen, J.; Zhao, T.; Fan, Z.
Granzyme K degrades the redox/DNA repair enzyme Ape1 to trigger oxidative stress of target cells leading to cytotoxicity
Mol. Immunol.
45
2225-2235
2008
Homo sapiens
brenda
Bovenschen, N.; Quadir, R.; van den Berg, A.L.; Brenkman, A.B.; Vandenberghe, I.; Devreese, B.; Joore, J.; Kummer, J.A.
Granzyme K displays highly restricted substrate specificity that only partially overlaps with granzyme A
J. Biol. Chem.
284
3504-3512
2009
Homo sapiens (P49863)
brenda
Bratke, K.; Klug, A.; Julius, P.; Kuepper, M.; Lommatzsch, M.; Sparmann, G.; Luttmann, W.; Virchow, J.C.
Granzyme K: a novel mediator in acute airway inflammation
Thorax
63
1006-1011
2008
Homo sapiens (P49863), Homo sapiens
brenda
Jenkins, M.R.; Trapani, J.A.; Doherty, P.C.; Turner, S.J.
Granzyme K expressing cytotoxic T lymphocytes protects against influenza virus in granzyme AB-/- mice
Viral Immunol.
21
341-346
2008
Mus musculus (O35205)
brenda
Guo, Y.; Chen, J.; Shi, L.; Fan, Z.
Valosin-containing protein cleavage by granzyme K accelerates an endoplasmic reticulum stress leading to caspase-independent cytotoxicity of target tumor cells
J. Immunol.
185
5348-5359
2010
Homo sapiens
brenda
Cooper, D.M.; Pechkovsky, D.V.; Hackett, T.L.; Knight, D.A.; Granville, D.J.
Granzyme K activates protease-activated receptor-1
PLoS ONE
6
e21484
2011
Homo sapiens
brenda
Plasman, K.; Demol, H.; Bird, P.I.; Gevaert, K.; Van Damme, P.
Substrate specificities of the granzyme tryptases A and K
J. Proteome Res.
13
6067-6077
2014
Mus musculus, Homo sapiens (P49863), Homo sapiens
brenda
Wensink, A.C.; Kemp, V.; Fermie, J.; Garcia Laorden, M.I.; van der Poll, T.; Hack, C.E.; Bovenschen, N.
Granzyme K synergistically potentiates LPS-induced cytokine responses in human monocytes
Proc. Natl. Acad. Sci. USA
111
5974-5979
2014
Homo sapiens (P49863), Homo sapiens
brenda
Dutta, D.; Park, I.; Guililat, H.; Sang, S.; Talapatra, A.; Singhal, B.; Mills, N.C.
Testosterone regulates granzyme K expression in rat testes
Endocr. Regul.
51
193-204
2017
Rattus norvegicus (P49864)
brenda
Sharma, M.; Merkulova, Y.; Raithatha, S.; Parkinson, L.G.; Shen, Y.; Cooper, D.; Granville, D.J.
Extracellular granzyme K mediates endothelial activation through the cleavage of protease-activated receptor-1
FEBS J.
283
1734-1747
2016
Homo sapiens (P49863), Homo sapiens
brenda
Turner, C.T.; Zeglinski, M.R.; Richardson, K.C.; Zhao, H.; Shen, Y.; Papp, A.; Bird, P.I.; Granville, D.J.
Granzyme K expressed by classically activated macrophages contributes to inflammation and impaired remodeling
J. Invest. Dermatol.
139
930-939
2019
Mus musculus (O35205), Homo sapiens (P49863), Homo sapiens
brenda