Calpains — An elaborate proteolytic system

Biochimica et Biophysica Acta (BBA) - Proteins and Proteomics - Tập 1824 Số 1 - Trang 224-236 - 2012
Yasuko Ono1, Hiroyuki Sorimachi
1Calpain Project, Department of Advanced Science for Biomolecules, Tokyo Metropolitan Institute of medical Science, Tokyo, Japan. [email protected]

Tóm tắt

Từ khóa


Tài liệu tham khảo

Croall, 1991, Calcium-activated neutral protease (calpain) system: structure, function, and regulation, Physiol. Rev., 71, 813, 10.1152/physrev.1991.71.3.813

Goll, 2003, The calpain system, Physiol. Rev., 83, 731, 10.1152/physrev.00029.2002

Suzuki, 2004, Structure, activation, and biology of calpain, Diabetes, 53, S12, 10.2337/diabetes.53.2007.S12

Sorimachi, 2011, Calpain chronicle — an enzyme family under multidisciplinary characterization, Proc. Jpn. Acad. Ser. B Phys. Biol. Sci., 87, 287, 10.2183/pjab.87.287

Sorimachi, 2011, Impact of genetic insights into calpain biology, J. Biochem., 150, 23, 10.1093/jb/mvr070

Guroff, 1964, A neutral calcium-activated proteinase from the soluble fraction of rat brain, J. Biol. Chem., 239, 149, 10.1016/S0021-9258(18)51762-2

Huston, 1968, Activation of skeletal muscle phosphorylase kinase by Ca2+. II. Identification of the kinase activating factor as a proteolytic enzyme, Biochemistry, 7, 2116, 10.1021/bi00846a014

Busch, 1972, Ca2+-specific removal of Z lines from rabbit skeletal muscle, J. Cell Biol., 52, 367, 10.1083/jcb.52.2.367

Meyer, 1964, Activation of skeletal muscle phosphorylase kinase by Ca2+, Biochemistry, 3, 1033, 10.1021/bi00896a004

Ishiura, 1978, Studies of a calcium-activated neutral protease from chicken skeletal muscle. I. Purification and characterization, J. Biochem., 84, 225, 10.1093/oxfordjournals.jbchem.a132111

Marchler-Bauer, 2009, CDD: specific functional annotation with the Conserved Domain Database, Nucleic Acids Res., 37, D205, 10.1093/nar/gkn845

Carragher, 2006, Calpain inhibition: a therapeutic strategy targeting multiple disease states, Curr. Pharm. Des., 12, 615, 10.2174/138161206775474314

Biswas, 2004, Calpains: targets of cataract prevention?, Trends Mol. Med., 10, 78, 10.1016/j.molmed.2003.12.007

Branca, 2004, Calpain-related diseases, Biochem. Biophys. Res. Commun., 322, 1098, 10.1016/j.bbrc.2004.07.126

Yamashima, 2004, Ca2+-dependent proteases in ischemic neuronal death: a conserved ‘calpain–cathepsin cascade’ from nematodes to primates, Cell Calcium, 36, 285, 10.1016/j.ceca.2004.03.001

Badalamente, 2000, Delay of muscle degeneration and necrosis in mdx mice by calpain inhibition, Muscle Nerve, 23, 106, 10.1002/(SICI)1097-4598(200001)23:1<106::AID-MUS14>3.0.CO;2-D

Bartus, 1994, Calpain inhibitor AK295 protects neurons from focal brain ischemia. Effects of postocclusion intra-arterial administration, Stroke, 25, 2265, 10.1161/01.STR.25.11.2265

Sugita, 1980, Ca-activated neutral protease and its inhibitors: in vitro effect on intact myofibrils, Muscle Nerve, 3, 335, 10.1002/mus.880030410

Sorimachi, 2010, Expanding members and roles of the calpain superfamily and their genetically modified animals, Exp. Anim., 59, 549, 10.1538/expanim.59.549

Tanaka, 2009, The proteasome: overview of structure and functions, Proc. Jpn. Acad. Ser. B Phys. Biol. Sci., 85, 12, 10.2183/pjab.85.12

Mizushima, 2010, Autophagy in mammalian development and differentiation, Nat. Cell Biol., 12, 823, 10.1038/ncb0910-823

Tait, 2010, Mitochondria and cell death: outer membrane permeabilization and beyond, Nat. Rev. Mol. Cell. Biol., 11, 621, 10.1038/nrm2952

Hosfield, 1999, Crystal structure of calpain reveals the structural basis for Ca2+-dependent protease activity and a novel mode of enzyme activation, EMBO J., 18, 6880, 10.1093/emboj/18.24.6880

Strobl, 2000, The crystal structure of calcium-free human m-calpain suggests an electrostatic switch mechanism for activation by calcium, Proc. Natl. Acad. Sci. U.S.A., 97, 588, 10.1073/pnas.97.2.588

Moldoveanu, 2002, A Ca2+ switch aligns the active site of calpain, Cell, 108, 649, 10.1016/S0092-8674(02)00659-1

Moldoveanu, 2003, Calpain silencing by a reversible intrinsic mechanism, Nat. Struct. Biol., 10, 371, 10.1038/nsb917

Moldoveanu, 2008, Concerted multi-pronged attack by calpastatin to occlude the catalytic cleft of heterodimeric calpains, Nature, 456, 404, 10.1038/nature07353

Hanna, 2008, Calcium-bound structure of calpain and its mechanism of inhibition by calpastatin, Nature, 456, 409, 10.1038/nature07451

Suzuki, 1995, Calpain: novel family members, activation, and physiologic function, Biol. Chem. Hoppe Seyler, 376, 523

Ohno, 1984, Evolutionary origin of a calcium-dependent protease by fusion of genes for a thiol protease and a calcium-binding protein?, Nature, 312, 566, 10.1038/312566a0

Maki, 1997, A growing family of the Ca2+-binding proteins with five EF-hand motifs, Biochem. J., 328, 718

Ono, 1998, Structure and physiology of calpain, an enigmatic protease, Biochem. Biophys. Res. Commun., 245, 289, 10.1006/bbrc.1998.8085

Sorimachi, 1993, A novel tissue-specific calpain species expressed predominantly in the stomach comprises two alternative splicing products with and without Ca2+-binding domain, J. Biol. Chem., 268, 19476, 10.1016/S0021-9258(19)36540-8

Yorikawa, 2008, Human calpain 7/PalBH associates with a subset of ESCRT-III-related proteins in its N-terminal region and partly localizes to endocytic membrane compartments, J. Biochem., 143, 731, 10.1093/jb/mvn030

Osako, 2010, Autolytic activity of human calpain 7 is enhanced by ESCRT-III-related protein IST1 through MIT-MIM interaction, FEBS J., 277, 4412, 10.1111/j.1742-4658.2010.07822.x

Ojima, 2011, Non-proteolytic functions of calpain-3 in sarcoplasmic reticulum in skeletal muscles, J. Mol. Biol., 407, 439, 10.1016/j.jmb.2011.01.057

Macqueen, 2010, A newly classified vertebrate calpain protease, directly ancestral to CAPN1 and 2, episodically evolved a restricted physiological function in placental mammals, Mol. Biol. Evol., 27, 1886, 10.1093/molbev/msq071

Macqueen, 2010, Characterisation of capn1, capn2-like, capn3 and capn11 genes in Atlantic halibut (Hippoglossus hippoglossus L.): transcriptional regulation across tissues and in skeletal muscle at distinct nutritional states, Gene, 453, 45, 10.1016/j.gene.2010.01.002

Andresen, 1991, Characterization of cDNA clones encoding a novel calcium-activated neutral proteinase from Schistosoma mansoni, J. Biol. Chem., 266, 15085, 10.1016/S0021-9258(18)98590-X

Friedrich, 2004, The calpain-system of Drosophila melanogaster: coming of age, Bioessays, 26, 1088, 10.1002/bies.20106

Syntichaki, 2002, Specific aspartyl and calpain proteases are required for neurodegeneration in C. elegans, Nature, 419, 939, 10.1038/nature01108

Ersfeld, 2005, Evolutionary relationships and protein domain architecture in an expanded calpain superfamily in kinetoplastid parasites, J. Mol. Evol., 61, 742, 10.1007/s00239-004-0272-8

Lid, 2002, The defective kernel 1 (dek1) gene required for aleurone cell development in the endosperm of maize grains encodes a membrane protein of the calpain gene superfamily, Proc. Natl. Acad. Sci. U.S.A., 99, 5460, 10.1073/pnas.042098799

Correa, 2001, Identification, classification and expression pattern analysis of sugarcane cysteine proteinases, Genet. Mol. Biol., 24, 275, 10.1590/S1415-47572001000100036

Futai, 1999, Aspergillus oryzae palBory encodes a calpain-like protease: homology to Emericella nidulans PalB and conservation of functional regions, J. Biosci. Bioeng., 88, 438, 10.1016/S1389-1723(99)80223-0

Dear, 1999, CAPN11: a calpain with high mRNA levels in testis and located on chromosome 6, Genomics, 59, 243, 10.1006/geno.1999.5859

Hata, 2010, Calpain 8/nCL-2 and calpain 9/nCL-4 constitute an active protease complex, G-calpain, involved in gastric mucosal defense, PLoS Genet., 6, e1001040, 10.1371/journal.pgen.1001040

Cao, 2001, xCL-2 is a novel m-type calpain and disrupts morphogenetic movements during embryogenesis in Xenopus laevis, Dev. Growth Differ., 43, 563, 10.1046/j.1440-169X.2001.00592.x

Futai, 2001, Molecular cloning of PalBH, a mammalian homologue of the Aspergillus atypical calpain PalB, Biochim. Biophys. Acta, 1517, 316, 10.1016/S0167-4781(00)00256-6

Caddick, 1986, Regulation of gene expression by pH of the growth medium in Aspergillus nidulans, Mol. Gen. Genet., 203, 346, 10.1007/BF00333978

Denison, 1995, Signaling of ambient pH in Aspergillus involves a cysteine protease, J. Biol. Chem., 270, 28519, 10.1074/jbc.270.48.28519

Negrete-Urtasun, 1999, Ambient pH signal transduction in Aspergillus: completion of gene characterization, Mol. Microbiol., 33, 994, 10.1046/j.1365-2958.1999.01540.x

Futai, 1999, The protease activity of a calpain-like cysteine protease in Saccharomyces cerevisiae is required for alkaline adaptation and sporulation, Mol. Gen. Genet., 260, 559, 10.1007/s004380050929

Hayashi, 2005, Constitutive activation of the pH-responsive Rim101 pathway in yeast mutants defective in late steps of the MVB/ESCRT pathway, Mol. Cell. Biol., 25, 9478, 10.1128/MCB.25.21.9478-9490.2005

Barnes, 1996, The tra-3 sex determination gene of Caenorhabditis elegans encodes a member of the calpain regulatory protease family, EMBO J., 15, 4477, 10.1002/j.1460-2075.1996.tb00825.x

Mugita, 1997, Identification of a novel, tissue-specific calpain htra-3; a human homologue of the Caenorhabditis elegans sex determination gene, Biochem. Biophys. Res. Commun., 239, 845, 10.1006/bbrc.1997.7571

Delaney, 1991, Molecular cloning and analysis of small optic lobes, a structural brain gene of Drosophila melanogaster, Proc. Natl. Acad. Sci. U.S.A., 88, 7214, 10.1073/pnas.88.16.7214

Kamei, 1998, SOLH, a human homologue of the Drosophila melanogaster small optic lobes gene is a member of the calpain and zinc-finger gene families and maps to human chromosome 16p13.3 near CATM (cataract with microphthalmia), Genomics, 51, 197, 10.1006/geno.1998.5395

Kamei, 2000, Solh, the mouse homologue of the Drosophila melanogaster small optic lobes gene: organization, chromosomal mapping, and localization of gene product to the olfactory bulb, Genomics, 64, 82, 10.1006/geno.1999.6098

Wang, 2003, The calpain domain of the maize DEK1 protein contains the conserved catalytic triad and functions as a cysteine proteinase, J. Biol. Chem., 278, 34467, 10.1074/jbc.M300745200

Margis, 2003, Phytocalpains: orthologous calcium-dependent cysteine proteinases, Trends Plant Sci., 8, 58, 10.1016/S1360-1385(02)00011-0

Ahn, 2004, Phytocalpain controls the proliferation and differentiation fates of cells in plant organ development, Plant J., 38, 969, 10.1111/j.1365-313X.2004.02102.x

Hibara, 2009, The ADAXIALIZED LEAF1 gene functions in leaf and embryonic pattern formation in rice, Dev. Biol., 334, 345, 10.1016/j.ydbio.2009.07.042

Johnson, 2005, AtDEK1 is essential for specification of embryonic epidermal cell fate, Plant J., 44, 114, 10.1111/j.1365-313X.2005.02514.x

Lid, 2005, Mutation in the Arabidopisis thaliana DEK1 calpain gene perturbs endosperm and embryo development while over-expression affects organ development globally, Planta, 221, 339, 10.1007/s00425-004-1448-6

Johnson, 2008, The phytocalpain defective kernel 1 is a novel Arabidopsis growth regulator whose activity is regulated by proteolytic processing, Plant Cell, 20, 2619, 10.1105/tpc.108.059964

Tian, 2007, Subcellular localization and functional domain studies of DEFECTIVE KERNEL1 in maize and Arabidopsis suggest a model for aleurone cell fate specification involving CRINKLY4 and SUPERNUMERARY ALEURONE LAYER1, Plant Cell, 19, 3127, 10.1105/tpc.106.048868

Dutt, 2006, m-Calpain is required for preimplantation embryonic development in mice, BMC Dev. Biol., 6, 3, 10.1186/1471-213X-6-3

Richard, 1995, Mutations in the proteolytic enzyme calpain 3 cause limb-girdle muscular dystrophy type 2A, Cell, 81, 27, 10.1016/0092-8674(95)90368-2

Yamada, 2009, Inhibition of calpain increases LIS1 expression and partially rescues in vivo phenotypes in a mouse model of lissencephaly, Nat. Med., 15, 1202, 10.1038/nm.2023

Yamada, 2010, A novel strategy for therapeutic intervention for the genetic disease: preventing proteolytic cleavage using small chemical compound, Int. J. Biochem. Cell Biol., 42, 1401, 10.1016/j.biocel.2010.05.017

Sorimachi, 1989, Molecular cloning of a novel mammalian calcium-dependent protease distinct from both m- and mu-types. Specific expression of the mRNA in skeletal muscle, J. Biol. Chem., 264, 20106, 10.1016/S0021-9258(19)47225-6

Dear, 1999, Diverse mRNA expression patterns of the mouse calpain genes Capn5, Capn6 and Capn11 during development, Mech. Dev., 89, 201, 10.1016/S0925-4773(99)00214-2

Dear, 1997, A new subfamily of vertebrate calpains lacking a calmodulin-like domain: implications for calpain regulation and evolution, Genomics, 45, 175, 10.1006/geno.1997.4870

Dear, 2000, Gene structure, chromosomal localization, and expression pattern of Capn12, a new member of the calpain large subunit gene family, Genomics, 68, 152, 10.1006/geno.2000.6289

Sorimachi, 1996, Highly conserved structure in the promoter region of the gene for muscle-specific calpain, p94, Biol. Chem., 377, 859

Kawabata, 2003, Newly identified exons encoding novel variants of p94/calpain 3 are expressed ubiquitously and overlap the α-glucosidase C gene, FEBS Lett., 555, 623, 10.1016/S0014-5793(03)01324-3

Hata, 2001, Both the conserved and the unique gene structure of stomach-specific calpains reveal processes of calpain gene evolution, J. Mol. Evol., 53, 191, 10.1007/s002390010209

Imajoh, 1987, The COOH-terminal E-F hand structure of calcium-activated neutral protease (CANP) is important for the association of subunits and resulting proteolytic activity, J. Biochem., 101, 447, 10.1093/oxfordjournals.jbchem.a121930

Lin, 1997, Crystal structure of calcium bound domain VI of calpain at 1.9Å resolution and its role in enzyme assembly, regulation, and inhibitor binding, Nat. Struct. Biol., 4, 539, 10.1038/nsb0797-539

Blanchard, 1997, Structure of a calpain Ca2+-binding domain reveals a novel EF-hand and Ca2+-induced conformational changes, Nat. Struct. Biol., 4, 532, 10.1038/nsb0797-532

Yoshizawa, 1995, A catalytic subunit of calpain possesses full proteolytic activity, FEBS Lett., 358, 101, 10.1016/0014-5793(94)01401-L

Arthur, 2000, Disruption of the murine calpain small subunit gene, Capn4: calpain is essential for embryonic development but not for cell growth and division, Mol. Cell. Biol., 20, 4474, 10.1128/MCB.20.12.4474-4481.2000

Zimmerman, 2000, The calpain small subunit gene is essential: its inactivation results in embryonic lethality, IUBMB Life, 50, 63, 10.1080/15216540050176610

Maki, 2002, Structures, functions and molecular evolution of the penta-EF-hand Ca2+-binding proteins, Biochim. Biophys. Acta, 1600, 51, 10.1016/S1570-9639(02)00444-2

Schad, 2002, A novel human small subunit of calpains, Biochem. J., 362, 383, 10.1042/0264-6021:3620383

Kiss, 2008, Local structural preferences of calpastatin, the intrinsically unstructured protein inhibitor of calpain, Biochemistry, 47, 6936, 10.1021/bi800201a

Hata, 2007, Stomach-specific calpain, nCL-2/calpain 8, is active without calpain regulatory subunit and oligomerizes through C2-like domains, J. Biol. Chem., 282, 27847, 10.1074/jbc.M703168200

Lee, 1999, Characterization of a human digestive tract-specific calpain, nCL-4, expressed in the baculovirus system, Arch. Biochem. Biophys., 362, 22, 10.1006/abbi.1998.1021

Ono, 2004, Possible regulation of the conventional calpain system by skeletal muscle-specific calpain, p94/calpain 3, J. Biol. Chem., 279, 2761, 10.1074/jbc.M308789200

Emori, 1987, Endogenous inhibitor for calcium-dependent cysteine protease contains four internal repeats that could be responsible for its multiple reactive sites, Proc. Natl. Acad. Sci. U.S.A., 84, 3590, 10.1073/pnas.84.11.3590

Maki, 1987, Repetitive region of calpastatin is a functional unit of the proteinase inhibitor, Biochem. Biophys. Res. Commun., 143, 300, 10.1016/0006-291X(87)90665-6

Maki, 1987, All four internally repetitive domains of pig calpastatin possess inhibitory activities against calpains I and II, FEBS Lett., 223, 174, 10.1016/0014-5793(87)80531-8

Pontremoli, 1988, The role of calpain and protein kinase C in activation of human neutrophils, Prog. Clin. Biol. Res., 282, 195

Noguchi, 1997, Functional cleavage of the common cytokine receptor γ chain (γc) by calpain, Proc. Natl. Acad. Sci. U.S.A., 94, 11534, 10.1073/pnas.94.21.11534

Nakagawa, 2000, Cross-talk between two cysteine protease families. Activation of caspase-12 by calpain in apoptosis, J. Cell Biol., 150, 887, 10.1083/jcb.150.4.887

Danial, 2004, Cell death: critical control points, Cell, 116, 205, 10.1016/S0092-8674(04)00046-7

Wells, 2005, Calpain proteases in cell adhesion and motility, Int. Rev. Cytol., 245, 1, 10.1016/S0074-7696(05)45001-9

Mellgren, 2007, Calpain is required for the rapid, calcium-dependent repair of wounded plasma membrane, J. Biol. Chem., 282, 2567, 10.1074/jbc.M604560200

Leloup, 2010, m-Calpain activation is regulated by its membrane localization and by its binding to phosphatidylinositol 4,5-bisphosphate, J. Biol. Chem., 285, 33549, 10.1074/jbc.M110.123604

Diez, 2002, Activation of the Aspergillus PacC zinc finger transcription factor requires two proteolytic steps, EMBO J., 21, 1350, 10.1093/emboj/21.6.1350

Hervas-Aguilar, 2007, Evidence for the direct involvement of the proteasome in the proteolytic processing of the Aspergillus nidulans zinc finger transcription factor PacC, J. Biol. Chem., 282, 34735, 10.1074/jbc.M706723200

Su, 1993, Molecular characterization of the yeast meiotic regulatory gene RIM1, Nucleic Acids Res., 21, 3789, 10.1093/nar/21.16.3789

Li, 1997, Proteolytic activation of Rim1p, a positive regulator of yeast sporulation and invasive growth, Genetics, 145, 63, 10.1093/genetics/145.1.63

Blanchin-Roland, 2005, ESCRT-I components of the endocytic machinery are required for Rim101-dependent ambient pH regulation in the yeast Yarrowia lipolytica, Microbiology, 151, 3627, 10.1099/mic.0.28196-0

Rodriguez-Galan, 2009, Physiological involvement in pH signaling of Vps24-mediated recruitment of Aspergillus PalB cysteine protease to ESCRT-III, J. Biol. Chem., 284, 4404, 10.1074/jbc.M808645200

Kullas, 2004, Snf7p, a component of the ESCRT-III protein complex, is an upstream member of the RIM101 pathway in Candida albicans, Eukaryot. Cell, 3, 1609, 10.1128/EC.3.6.1609-1618.2004

Xu, 2004, Multivesicular body-ESCRT components function in pH response regulation in Saccharomyces cerevisiae and Candida albicans, Mol. Biol. Cell, 15, 5528, 10.1091/mbc.E04-08-0666

Missotten, 1999, Alix, a novel mouse protein undergoing calcium-dependent interaction with the apoptosis-linked-gene 2 (ALG-2) protein, Cell Death Differ., 6, 124, 10.1038/sj.cdd.4400456

Vito, 1999, Cloning of AIP1, a novel protein that associates with the apoptosis-linked gene ALG-2 in a Ca2+-dependent reaction, J. Biol. Chem., 274, 1533, 10.1074/jbc.274.3.1533

Katoh, 2003, The ALG-2-interacting protein Alix associates with CHMP4b, a human homologue of yeast Snf7 that is involved in multivesicular body sorting, J. Biol. Chem., 278, 39104, 10.1074/jbc.M301604200

Sadoul, 2006, Do Alix and ALG-2 really control endosomes for better or for worse?, Biol. Cell, 98, 69, 10.1042/BC20050007

Richard, 2000, Loss of calpain 3 proteolytic activity leads to muscular dystrophy and to apoptosis-associated IκBα/nuclear factor κB pathway perturbation in mice, J. Cell Biol., 151, 1583, 10.1083/jcb.151.7.1583

Kramerova, 2004, Null mutation of calpain 3 (p94) in mice causes abnormal sarcomere formation in vivo and in vitro, Hum. Mol. Genet., 13, 1373, 10.1093/hmg/ddh153

Ojima, 2010, Dynamic distribution of muscle-specific calpain in mice has a key role in physical-stress adaptation and is impaired in muscular dystrophy, J. Clin. Invest., 120, 2672, 10.1172/JCI40658

Solomon, 1998, The N-end rule pathway catalyzes a major fraction of the protein degradation in skeletal muscle, J. Biol. Chem., 273, 25216, 10.1074/jbc.273.39.25216

Huang, 1998, Role of calpain in skeletal-muscle protein degradation, Proc. Natl. Acad. Sci. U.S.A., 95, 12100, 10.1073/pnas.95.21.12100

Sorimachi, 1993, Muscle-specific calpain, p94, is degraded by autolysis immediately after translation, resulting in disappearance from muscle, J. Biol. Chem., 268, 10593, 10.1016/S0021-9258(18)82240-2

Ono, 2010, Skeletal muscle-specific calpain is an intracellular Na+-dependent protease, J. Biol. Chem., 285, 22986, 10.1074/jbc.M110.126946

Hayashi, 2008, Multiple molecular interactions implicate the connectin/titin N2A region as a modulating scaffold for p94/calpain 3 activity in skeletal muscle, J. Biol. Chem., 283, 14801, 10.1074/jbc.M708262200

Sorimachi, 1995, Muscle-specific calpain, p94, responsible for limb girdle muscular dystrophy type 2A, associates with connectin through IS2, a p94-specific sequence, J. Biol. Chem., 270, 31158, 10.1074/jbc.270.52.31158

Beckmann, 2008, Calpain 3, the “gatekeeper” of proper sarcomere assembly, turnover and maintenance, Neuromuscul. Disord., 18, 913, 10.1016/j.nmd.2008.08.005

Ojima, 2007, Myogenic stage, sarcomere length, and protease activity modulate localization of muscle-specific calpain, J. Biol. Chem., 282, 14493, 10.1074/jbc.M610806200

Hata, 2006, Stomach-specific calpain, nCL-2, localizes in mucus cells and proteolyzes the β-subunit of coatomer complex, β-COP, J. Biol. Chem., 281, 11214, 10.1074/jbc.M509244200

Hata, 2001, Domain II of m-calpain is a Ca2+-dependent cysteine protease, FEBS Lett., 501, 111, 10.1016/S0014-5793(01)02611-4

Rey, 2002, The protease core of the muscle-specific calpain, p94, undergoes Ca2+-dependent intramolecular autolysis, FEBS Lett., 532, 401, 10.1016/S0014-5793(02)03722-5

Davis, 2007, The crystal structures of human calpains 1 and 9 imply diverse mechanisms of action and auto-inhibition, J. Mol. Biol., 366, 216, 10.1016/j.jmb.2006.11.037

Moldoveanu, 2004, Calpain activation by cooperative Ca2+ binding at two non-EF-hand sites, J. Biol. Chem., 279, 6106, 10.1074/jbc.M310460200

Yuan, 2010, Structure of the human BK channel Ca2+-activation apparatus at 3.0Å resolution, Science, 329, 182, 10.1126/science.1190414

Moldoveanu, 2004, Crystal structures of calpain-E64 and -leupeptin inhibitor complexes reveal mobile loops gating the active site, J. Mol. Biol., 343, 1313, 10.1016/j.jmb.2004.09.016

Tsuji, 1981, Studies on a Ca2+-activated neutral proteinase of rabbit skeletal muscle. II. Characterization of sulfhydryl groups and a role of Ca2+ ions in this enzyme, J. Biochem., 90, 1405, 10.1093/oxfordjournals.jbchem.a133606

Dainese, 2002, Conformational changes of calpain from human erythrocytes in the presence of Ca2+, J. Biol. Chem., 277, 40296, 10.1074/jbc.M204471200

Herasse, 1999, Expression and functional characteristics of calpain 3 isoforms generated through tissue-specific transcriptional and posttranscriptional events, Mol. Cell. Biol., 19, 4047, 10.1128/MCB.19.6.4047

Saido, 1992, Positive regulation of mu-calpain action by polyphosphoinositides, J. Biol. Chem., 267, 24585, 10.1016/S0021-9258(18)35804-6

Tompa, 2001, Domain III of calpain is a Ca2+-regulated phospholipid-binding domain, Biochem. Biophys. Res. Commun., 280, 1333, 10.1006/bbrc.2001.4279

Shao, 2006, Spatial localization of m-calpain to the plasma membrane by phosphoinositide biphosphate binding during epidermal growth factor receptor-mediated activation, Mol. Cell. Biol., 26, 5481, 10.1128/MCB.02243-05

Saido, 1993, In situ capture of mu-calpain activation in platelets, J. Biol. Chem., 268, 7422, 10.1016/S0021-9258(18)53191-4

Saido, 1993, Spatial resolution of fodrin proteolysis in postischemic brain, J. Biol. Chem., 268, 25239, 10.1016/S0021-9258(19)74593-1

Horikawa, 2000, Genetic variation in the gene encoding calpain-10 is associated with type 2 diabetes mellitus, Nat. Genet., 26, 163, 10.1038/79876

Sokol, 2000, Proteolysis in Caenorhabditis elegans sex determination: cleavage of TRA-2A by TRA-3, Genes Dev., 14, 901, 10.1101/gad.14.8.901

Tonami, 2011, Calpain 6, a microtubule-stabilizing protein, regulates Rac1 activity and cell motility through interaction with GEF-H1, J. Cell Sci., 124, 1214, 10.1242/jcs.072561

Tonami, 2007, Calpain 6 is involved in microtubule stabilization and cytoskeletal organization, Mol. Cell. Biol., 27, 2548, 10.1128/MCB.00992-06

Stabach, 1997, Site-directed mutagenesis of alpha II spectrin at codon 1175 modulates its mu-calpain susceptibility, Biochemistry, 36, 57, 10.1021/bi962034i

Tompa, 2004, On the sequential determinants of calpain cleavage, J. Biol. Chem., 279, 20775, 10.1074/jbc.M313873200

Cuerrier, 2005, Determination of peptide substrate specificity for mu-calpain by a peptide library-based approach: the importance of primed side interactions, J. Biol. Chem., 280, 40632, 10.1074/jbc.M506870200

duVerle, 2010, CaMPDB: a resource for calpain and modulatory proteolysis, Genome Inform., 22, 202, 10.1142/9781848165786_0017

duVerle, 2011, Calpain cleavage prediction using multiple kernel learning, PLoS One, 6, e19035, 10.1371/journal.pone.0019035

Backes, 2005, GraBCas: a bioinformatics tool for score-based prediction of Caspase- and Granzyme B-cleavage sites in protein sequences, Nucleic Acids Res., 33, W208, 10.1093/nar/gki433

Yang, 2005, Prediction of caspase cleavage sites using Bayesian bio-basis function neural networks, Bioinformatics, 21, 1831, 10.1093/bioinformatics/bti281

Song, 2010, Cascleave: towards more accurate prediction of caspase substrate cleavage sites, Bioinformatics, 26, 752, 10.1093/bioinformatics/btq043

Holm, 1993, Protein structure comparison by alignment of distance matrices, J. Mol. Biol., 233, 123, 10.1006/jmbi.1993.1489