Cysteine cathepsins and caspases in silicosis

Biological Chemistry

Volumn 387, Issue 7, 2006, Pages 863-870

  Lalmanach, Gilles ^a   Diot, Elisabeth ^a   Godat, Emmanuel ^a   Lecaille, Fabien ^a   Hervé Grépinet, Virginie ^a  

^a UNIVERSITÉ DE TOURS   (France)

Author keywords

Apoptosis; Bronchoalveolar lavage; Fibrosis; Inflammation; Protease; Protease inhibitor

Indexed keywords

ALPHA 1 ANTITRYPSIN; CASPASE; CATHEPSIN; COLLAGEN; CYSTEINE; FAS ANTIGEN; FAS LIGAND; MATRIX METALLOPROTEINASE; PROTEINASE INHIBITOR; SECRETORY LEUKOCYTE PROTEINASE INHIBITOR; SILICON DIOXIDE; TISSUE INHIBITOR OF METALLOPROTEINASE;

APOPTOSIS; BASEMENT MEMBRANE; CATALYSIS; CELL DEATH; CONFERENCE PAPER; CYSTIC FIBROSIS; EMPHYSEMA; ENZYME ACTIVITY; ENZYME DEGRADATION; ENZYME INACTIVATION; ENZYME INHIBITION; ENZYME RELEASE; EXTRACELLULAR MATRIX; HUMAN; INFLAMMATION; LUNG ALVEOLUS MACROPHAGE; LUNG LAVAGE; NONHUMAN; OCCUPATIONAL DISEASE; OCCUPATIONAL EXPOSURE; OCULAR ALBINISM; PATHOGENESIS; PHAGOCYTE; PNEUMOCONIOSIS; PRIORITY JOURNAL; PROTEIN EXPRESSION; PROTEIN SYNTHESIS; SILICOSIS;

ANTIGENS, CD95; APOPTOSIS; BRONCHOALVEOLAR LAVAGE FLUID; CASPASES; CATHEPSINS; CYSTEINE; CYSTEINE PROTEINASE INHIBITORS; FAS LIGAND PROTEIN; HUMANS; MEMBRANE GLYCOPROTEINS; SILICOSIS; TUMOR NECROSIS FACTORS;

EID: 33746192084     PISSN: 14316730     EISSN: 14374315     Source Type: Journal    
DOI: 10.1515/BC.2006.109     Document Type: Conference Paper

References (92)

1. Abrahamson, M., Buttle, D.J., Mason, R.W., Hansson, H., Grubb, A., Lilja, H., and Ohlsson, K. (1991a). Regulation of cystatin C activity by serine proteinases. Biomed. Biochim. Acta 50, 587-593.
2. Abrahamson, M., Mason, R.W., Hansson, H., Buttle, D.J., Grubb, A., and Ohlsson, K. (1991b). Human cystatin C. role of the N-terminal segment in the inhibition of human cysteine proteinases and in its inactivation by leucocyte elastase. Biochem. J. 273, 621-626.
3. Abrahamson, M., Alvarez-Fernandez, M., and Nathanson, C.M. (2003). Cystatins. Biochem. Soc. Symp. 70, 179-199.
4. Barnes, P.J. (1998). New therapies for chronic obstructive pulmonary disease. Thorax 53, 137-147.
5. Barnes, P.J., Shapiro, S.D., and Pauwels, R.A. (2003). Chronic obstructive pulmonary disease: molecular and cellular mechanisms. Eur. Respir. J. 22, 672-688.
6. Borges, V.M., Falcao, H., Leite-Junior, J.H., Alvim, L., Teixeira, G.P., Russo, M., Nobrega, A.F., Lopes, M.F., Rocco, P.M., Davidson, W.F., Linden, R., Yagita, H., Zin, W.A., and DosReis, G.A. (2001). Fas ligand triggers pulmonary silicosis. J. Exp. Med. 194, 155-164.
7. Borges, V.M., Lopes, M.F., Falcao, H., Leite-Junior, J.H., Rocco, P.R., Davidson, W.F., Linden, R., Zin, W.A., and DosReis, G.A. (2002). Apoptosis underlies immunopathogenic mechanisms in acute silicosis. Am. J. Respir. Cell. Mol. Biol. 27, 78-84.
8. Brain, J.D. (1986). Toxicological aspects of alterations of pulmonary macrophage function. Annu. Rev. Pharmacol. Toxicol. 26, 547-565.
9. Brasch, F., Ten Brinke, A., Johnen, G., Ochs, M., Kapp, N., Muller, K.M., Beers, M.F., Fehrenbach, H., Richter, J., Batenburg, J.J., and Buhling, F. (2002). Involvement of cathepsin H in the processing of the hydrophobic surfactant-associated protein C in type II pneumocytes. Am. J. Respir. Cell. Mol. Biol. 26, 659-670.
10. Buhling, F., Gerber, A., Ansorge, S., and Weite, T. (1999). Cathepsin cysteine proteinases in the lung. Pneumologie 53, 400-407.
11. Buhling, F., Reisenauer, A., Gerber, A., Kruger, S., Weber, E., Bromme, D., Roessner, A., Ansorge, S., Weite, T., and Rocken, C. (2001). Cathepsin K: a marker of macrophage differentiation? J. Pathol. 195, 375-382.
12. Buhling, F., Rocken, C., Brasch, F., Hartig, R., Yasuda, Y., Saftig, P., Bromme, D., and Weite, T. (2004). Pivotal role of cathepsin K in lung fibrosis. Am. J. Pathol. 164, 2203-2216.
13. Buhling, F., Wille, A., Rocken, C., Wiesner, O., Baier, A., Meinecke, I., Weite, T., and Pap, T. (2005). Altered expression of membrane-bound and soluble CD95/Fas contributes to the resistance of fibrotic lung fibroblasts to FasL induced apoptosis. Respir. Res. 6, 37.
14. Buttle, D.J., Abrahamson, M., Burnett, D., Mort, J.S., Barrett, A.J., Dando, P.M., and Hill, S.L. (1991). Human sputum cathepsin B degrades proteoglycan, is inhibited by α2-macroglobulin and is modulated by neutrophil elastase cleavage of cathepsin B precursor and cystatin C. Biochem. J. 276, 325-331.
15. Castranova, V. and Vallyathan, V. (2000). Silicosis and coal workers' pneumoconiosis. Environ. Health Perspect. 108 (Suppl. 4), 675-684.
16. Chang, J.C., Lesser, M., Yoo, O.H., and Orlowski, M. (1986). Increased cathepsin B-like activity in alveolar macrophages and bronchoalveolar lavage fluid from smokers. Am. Rev. Respir. Dis. 134, 538-541.
17. Chao, S.K., Hamilton, R.F., Pfau, J.C., and Holian, A. (2001). Cell surface regulation of silica-induced apoptosis by the SR-A scavenger receptor in a murine lung macrophage cell line (MH-S). Toxicol. Appl. Pharmacol. 174, 10-16.
18. Chapman, H.A. Jr. and Shi, G.P. (2000). Protease injury in the development of COPD. Chest 117, 295S-299S.
19. Chapman, H.A., Riese, R.J., and Shi, G.P. (1997). Emerging roles for cysteine proteases in human biology. Annu. Rev. Physiol. 59, 63-88.
20. Davidson, J.M. (1990). Biochemistry and turnover of lung interstitium. Eur. Respir. J. 3, 1048-1063.
21. Denault, J.B. and Salvesen, G.S. (2002). Caspases: keys in the ignition of cell death. Chem. Rev. 102, 4489-4500.
22. DosReis, G.A., Borges, V.M., and Zin, W.A. (2004). The central role of Fas-ligand cell signaling in inflammatory lung diseases. J. Cell. Mol. Med. 8, 285-293.
23. Everts, V., van der Zee, E., Creemers, L., and Beertsen, W. (1996). Phagocytosis and intracellular digestion of collagen, its role in turnover and remodelling. Histochem. J. 28, 229-245.
24. Feng, G.H., Bailin, T., Oh, J., and Spritz, R.A. (1997). Mouse pale ear (ep) is homologous to human Hermansky-Pudlak syndrome and contains a rare 'AT-AC' intron. Hum. Mol. Genet. 6, 793-697.
25. Ferri, K.F. and Kroemer, G. (2001). Organelle-specific initiation of cell death pathways. Nat. Cell Biol. 3, 255-263.
26. Fubini, B. and Hubbard, A. (2003). Reactive oxygen species (ROS) and reactive nitrogen species (RNS) generation by silica in inflammation and fibrosis. Free Radic. Biol. Med. 34, 1507-1516.
27. Fuentes-Prior, P. and Salvesen, G.S. (2004). The protein structures that shape caspase activity, specificity, activation and inhibition. Biochem. J. 384, 201-232.
28. Garnero, P., Borel, O., Byrjalsen, I., Ferreras, M., Drake, F.H., McQueney, M.S., Foged, N.T., Delmas, P.D., and Delaisse, J.M. (1998). The collagenolytic activity of cathepsin K is unique among mammalian proteinases. J. Biol. Chem. 273, 32347-32352.
29. Guicciardi, M.E., Deussing, J., Miyoshi, H., Bronk, S.F., Svingen, P.A., Peters, C., Kaufmann, S.H., and Gores, G.J. (2000). Cathepsin B contributes to TNF-α-mediated hepatocyte apoptosis by promoting mitochondrial release of cytochrome c. J. Clin. Invest. 106, 1127-1137.
30. Guttentag, S., Robinson, L., Zhang, P., Brasch, F., Buhling, F., and Beers, M. (2003). Cysteine protease activity is required for surfactant protein B processing and lamellar body genesis. Am. J. Respir. Cell. Mol. Biol. 28, 69-79.
31. Hannothiaux, M.H., Scharfman, A., Wastiaux, A., Cornu, L., van Brussel, E., Lafitte, J.J., Sebastien, P., and Roussel, P. (1991). An attempt to evaluate lung aggression in monkey silicosis: hydrolases, peroxidase and antiproteases activities in serial bronchoalveolar lavages. Eur. Respir. J. 4, 191-204.
32. Heidtmann, H.H., Salge, U., Havemann, K., Kirschke, H., and Wiederanders, B. (1993). Secretion of a latent, acid activatable cathepsin L precursor by human non-small cell lung cancer cell lines. Oncol. Res. 5, 441-451.
33. Henskens, Y.M., Veerman, E.C., and Nieuw Amerongen, A.V. (1996). Cystatins in health and disease. Biol. Chem. Hoppe-Seyler 377, 71-86.
34. Hermansky, F. and Pudlak, P. (1959). Albinism associated with hemorrhagic diathesis and unusual pigmented reticular cells in the bone marrow: report of two cases with histochemical studies. Blood 14, 162-169.
35. Ishii, Y., Hashizume, Y., Watanabe, T., Waguri, S., Sato, N., Yamamoto, M., Hasegawa, S., Kominami, E., and Uchiyama, Y. (1991). Cysteine proteinases in bronchoalveolar epithelial cells and lavage fluid of rat lung. J. Histochem. Cytochem. 39, 461-468.
36. Ivy, G.O., Kanai, S., Ohta, M., Smith, G., Sato, Y., Kobayashi, M., and Kitani, K. (1989). Lipofuscin-like substances accumulate rapidly in brain, retina and internal organs with cysteine protease inhibition. Adv. Exp. Med. Biol. 266, 31-45.
37. Iyer, R. and Holian, A. (1997). Involvement of the ICE family of proteases in silica-induced apoptosis in human alveolar macrophages. Am. J. Physiol. 273, L760-767.
38. Iyer, R., Hamilton, R.F., Li, L., and Holian, A. (1996). Silica-induced apoptosis mediated via scavenger receptor in human alveolar macrophages. Toxicol. Appl. Pharmacol. 141, 84-92.
39. Johnson, D.A., Barrett, A.J., and Mason, R.W. (1986). Cathepsin L inactivates a1-proteinase inhibitor by cleavage in the reactive site region. J. Biol. Chem. 261, 14748-14751.
40. Langley, R.J., Kalra, R., Mishra, N.C., Hahn, F.F., Razani-Boroujerdi, S., Singh, S.P., Benson, J.M., Pena-Philippides, J.C., Barr, E.B., and Sopori, M.L. (2004). A biphasic response to silica: I. Immunostimulation is restricted to the early stage of silicosis in Lewis rats. Am. J. Respir. Cell. Mol. Biol. 30, 823-829.
41. Lecaille, F., Kaleta, J., and Bromme, D. (2002). Human and parasitic papain-like cysteine proteases: their role in physiology and pathology and recent developments in inhibitor design. Chem. Rev. 102, 4459-4488.
42. Lesser, M., Padilla, M.L., and Cardozo, C. (1992). Induction of emphysema in hamsters by intratracheal instillation of cathepsin B. Am. Rev. Respir. Dis. 145, 661-668.
43. Luthgens, K., Ebert, W., Trefz, G., Gabrijelcic, D., Turk, V., and Lah, T. (1993). Cathepsin B and cysteine proteinase inhibitors in bronchoalveolar lavage fluid of lung cancer patients. Cancer Detect. Prev. 17, 387-397.
44. Majerle, A. and Jerala, R. (2003). Protein inhibitors form complexes with procathepsin L and augment cleavage of the propeptide. Arch. Biochem. Biophys. 417, 53-58.
45. Mossman, B.T. and Churg, A. (1998). Mechanisms in the pathogenesis of asbestosis and silicosis. Am. J. Respir. Crit. Care Med. 157, 1666-1680.
46. Oh, J., Bailin, T., Fukai, K., Feng, G.H., Ho, L., Mao, J.I., Frenk, E., Tamura, N., and Spritz, R.A. (1996). Positional cloning of a gene for Hermansky-Pudlak syndrome, a disorder of cytoplasmic organelles. Nat. Genet. 14, 300-306.
47. Ohashi, K., Naruto, M., Nakaki, T., and Sano, E. (2003). Identification of interleukin-8 converting enzyme as cathepsin L. Biochim. Biophys. Acta 1649, 30-39.
48. Owen, C.A. and Campbell, E.J. (1999a). The cell biology of leukocyte-mediated proteolysis. J. Leukoc. Biol. 65, 137-150.
49. Owen, C.A. and Campbell, E.J. (1999b). Extracellular proteolysis: new paradigms for an old paradox. J. Lab. Clin. Med. 134, 341-351.
50. Perdereau, C., Godat, E., Maurel, M.C., Hazouard, E., Diot, E., and Lalmanach, G. (2006). Cysteine cathepsins in human silicotic bronchoalveolar lavage fluids. Biochim. Biophys. Acta 1762, 351-356.
51. Perez-Ramos, J., de Lourdes Segura-Valdez, M., Vanda, B., Selman, M., and Pardo, A. (1999). Matrix metalloproteinases 2, 9, and 13, and tissue inhibitors of metalloproteinases 1 and 2 in experimental lung silicosis. Am. J. Respir. Crit. Care Med. 160, 1274-1282.
52. Pettus, B.J., Chalfant, C.E., and Hannun, Y.A. (2002). Ceramide in apoptosis: an overview and current perspectives. Biochim. Biophys. Acta 1585, 114-125.
53. Porter, D.W., Ye, J., Ma, J., Barger, M., Robinson, V.A., Ramsey, D., McLaurin, J., Khan, A., Landsittel, D., Teass, A., and Castranova, V. (2002). Time course of pulmonary response of rats to inhalation of crystalline silica: NF-κB activation, inflammation, cytokine production, and damage. Inhal. Toxicol. 14, 349-367.
54. Punturieri, A., Filippov, S., Allen, E., Caras, I., Murray, R., Reddy, V., and Weiss, S.J. (2000). Regulation of elastinolytic cysteine proteinase activity in normal and cathepsin K-deficient human macrophages. J. Exp. Med. 192, 789-799.
55. Reddy, V.Y., Zhang, Q.Y., and Weiss, S.J. (1995). Pericellular mobilization of the tissue-destructive cysteine proteinases, cathepsins B, L, and S, by human monocyte-derived macrophages. Proc. Natl. Acad. Sci. USA 92, 3849-3853.
56. Reilly, J.J. Jr., Chen, P., Sailor, L.Z., Wilcox, D., Mason, R.W., and Chapman, H.A. Jr. (1991). Cigarette smoking induces an elastolytic cysteine proteinase in macrophages distinct from cathepsin L. Am. J. Physiol. 261, L41-88.
57. Rosenman, K.D., Moore-Fuller, M., and Reilly, M.J. (1999). Connective tissue disease and silicosis. Am. J. Ind. Med. 35, 375-381.
58. Salvesen, G.S. and Abrams, J.M. (2004). Caspase activation-stepping on the gas or releasing the brakes? Lessons from humans and flies. Oncogene 23, 2774-2784.
59. Sarih, M., Souvannavong, V., Brown, S.C., and Adam, A. (1993). Silica induces apoptosis in macrophages and the release of interleukin-1α and interleukin-1β. J. Leukoc. Biol. 54, 407-413.
60. Scabilloni, J.F., Wang, L., Antonini, J.M., Roberts, J.R., Castranova, V., and Mercer, R.R. (2005). Matrix metalloproteinase induction in fibrosis and fibrotic nodule formation due to silica inhalation. Am. J. Physiol. Lung Cell. Mol. Physiol. 288, L709-L717.
61. Serveau-Avesque, C., Ferrer-Di Martino, M., Herve-Grepinet, V., Hazouard, E., Gauthier, F., Diot, E., and Lalmanach, G. (2006). Active cathepsins B, H, K, L and S in human inflammatory bronchoalveolar lavage fluids. Biol. Cell 98, 15-22.
62. Shangary, S., Lerner, E.C., Zhan, Q., Corey, S.J., Smithgall, T.E., and Baskaran, R. (2003). Lyn regulates the cell death response to ultraviolet radiation through c-Jun N terminal kinase-dependent Fas ligand activation. Exp. Cell. Res. 289, 67-76.
63. Shapiro, S.D. (2002). Proteinases in chronic obstructive pulmonary disease. Biochem. Soc. Trans. 30, 98-102.
64. Shen, H.M., Zhang, Z., Zhang, Q.F., and Ong, C.N. (2001). Reactive oxygen species and caspase activation mediate silica-induced apoptosis in alveolar macrophages. Am. J. Physiol. Lung Cell. Mol. Physiol. 280, L10-L17.
65. Shi, G.P., Bryant, R.A., Riese, R., Verhelst, S., Driessen, C., Li, Z., Bromme, D., Ploegh, H.L., and Chapman, H.A. (2000). Role for cathepsin F in invariant chain processing and major histocompatibility complex class II peptide loading by macrophages. J. Exp. Med. 191, 1177-1186.
66. Steenland, K. and Goldsmith, D.F. (1995). Silica exposure and autoimmune diseases. Am. J. Ind. Med. 28, 603-608.
67. Stockley, R.A. (1999). Neutrophils and protease/antiprotease imbalance. Am. J. Respir. Crit. Care Med. 160, S49-S52.
68. Stoka, V., Turk, B., Schendel, S.L., Kim, T.H., Cirman, T., Snipas, S.J., Ellerby, L.M., Bredesen, D., Freeze, H., Abrahamson, M., et al. (2001). Lysosomal protease pathways to apoptosis. Cleavage of bid, not pro-caspases, is the most likely route. J. Biol. Chem. 276, 3149-3157.
69. Taggart, C., Cervantes-Laurean, D., Kim, G., McElvaney, N.G., Wehr, N., Moss, J., and Levine, R.L. (2000). Oxidation of either methionine 351 or methionine 358 in α1-antitrypsin causes loss of anti-neutrophil elastase activity. J. Biol. Chem. 275, 27258-27265.
70. Taggart, C.C., Lowe, G.J., Greene, C.M., Mulgrew, A.T., O'Neill, S.J., Levine, R.L., and McElvaney, N.G. (2001). Cathepsin B, L, and S cleave and inactivate secretory leucoprotease inhibitor. J. Biol. Chem. 276, 33345-33352.
71. Taggart, C.C., Greene, C.M., Smith, S.G., Levine, R.L., McCray, P.B. Jr., O'Neill, S., and McElvaney, N.G. (2003). Inactivation of human b-defensins 2 and 3 by elastolytic cathepsins. J. Immunol. 171, 931-937.
72. Takahashi, H., Ishidoh, K., Muno, D., Ohwada, A., Nukiwa, T., Kominami, E., and Kira, S. (1993). Cathepsin L activity is increased in alveolar macrophages and bronchoalveolar lavage fluid of smokers. Am. Rev. Respir. Dis. 147, 1562-1568.
73. Takeyabu, K., Betsuyaku, T., Nishimura, M., Yoshioka, A., Tanino, M., Miyamoto, K., and Kawakami, Y. (1998). Cysteine proteinases and cystatin C in bronchoalveolar lavage fluid from subjects with subclinical emphysema. Eur. Respir. J. 12, 1033-1039.
74. Thibodeau, M., Giardina, C., and Hubbard, A.K. (2003). Silica-induced caspase activation in mouse alveolar macrophages is dependent upon mitochondrial integrity and aspartic proteolysis. Toxicol. Sci. 76, 91-101.
75. Thibodeau, M.S., Giardina, C., Knecht, D.A., Helble, J., and Hubbard, A.K. (2004). Silica-induced apoptosis in mouse alveolar macrophages is initiated by lysosomal enzyme activity. Toxicol. Sci. 80, 34-48.
76. Thome, M. and Tschopp, J. (2001). Regulation of lymphocyte proliferation and death by FLIP. Nat. Rev. Immunol. 1, 50-58.
77. Thornberry, N.A., Bull, H.G., Calaycay, J.R., Chapman, K.T., Howard, A.D., Kostura, M.J., Miller, D.K., Molineaux, S.M., Weidner, J.R., Aunins, J., et al. (1992). A novel heterodimeric cysteine protease is required for interleukin-1β processing in monocytes. Nature 356, 768-774.
78. Tomita, K., Caramori, G., Lim, S., Ito, K., Hanazawa, T., Oates, T., Chiselita, I., Jazrawi, E., Chung, K.F., Barnes, P.J., and Adcock, I.M. (2002). Increased p21(CIP1/WAF1) and B cell lymphoma leukemia-x(L) expression and reduced apoptosis in alveolar macrophages from smokers. Am. J. Respir. Crit. Care Med. 166, 724-731.
79. Travis, J. and Salvesen, G.S. (1983). Human plasma proteinase inhibitors. Annu. Rev. Biochem. 52, 655-709.
80. Turk, V. and Bode, W. (1991). The cystatins: protein inhibitors of cysteine proteinases. FEBS Lett. 285, 213-219.
81. Turk, B., Turk, D., and Turk, V. (2000). Lysosomal cysteine proteases: more than scavengers. Biochim. Biophys. Acta 1477, 98-111.
82. Turk, V., Turk, B., and Turk, D. (2001). Lysosomal cysteine proteases: facts and opportunities. EMBO J. 20, 4629-4633.
83. Ueki, A., Isozaki, Y., and Kusaka, M. (2005). Anti-caspase-8 autoantibody response in silicosis patients is associated with HLA-DRB1, DQB1 and DPB1 alleles. J. Occup. Health 47, 61-67.
84. Ulbricht, B., Hagmann, W., Ebert, W., and Spiess, E. (1996). Differential secretion of cathepsins B and L from normal and tumor human lung cells stimulated by 12(S)-hydroxy-eicosatetraenoic acid. Exp. Cell. Res. 226, 255-263.
85. Van den Brule, S., Misson, P., Buhling, F., Lison, D., and Huaux, F. (2005). Overexpression of cathepsin K during silica-induced lung fibrosis and control by TGF-β. Respir. Res. 6, 84.
86. Vanhee, D., Gosset, P., Boitelle, A., Wallaert, B., and Tonnel, A.B. (1995). Cytokines and cytokine network in silicosis and coal workers' pneumoconiosis. Eur. Respir. J. 8, 834-842.
87. Wang, Z., Zheng, T., Zhu, Z., Homer, R.J., Riese, R.J., Chapman, H.A. Jr., Shapiro, S.D., and Elias, J.A. (2000). Interferon γ induction of pulmonary emphysema in the adult murine lung. J. Exp. Med. 192, 1587-1600.
88. Wolters, P.J. and Chapman, H.A. (2000). Importance of lysosomal cysteine proteases in lung disease. Respir. Res. 1, 170-177.
89. Yasuda, Y., Li, Z., Greenbaum, D., Bogyo, M., Weber, E., and Bromme, D. (2004). Cathepsin V, a novel and potent elastolytic activity expressed in activated macrophages. J. Biol. Chem. 279, 36761-36770.
90. Yayoi, Y., Ohsawa, Y., Koike, M., Zhang, G., Kominami, E., and Uchiyama, Y. (2001). Specific localization of lysosomal aminopeptidases in type II alveolar epithelial cells of the rat lung. Arch. Histol. Cytol. 64, 89-97.
91. Yoshioka, Y., Kumasaka, T., Ishidoh, K., Kominami, E., Mitani, K., Hosokawa, Y., and Fukuchi, Y. (2004). Inflammatory response and cathepsins in silica-exposed Hermansky-Pudlak syndrome model pale ear mice. Pathol. Int. 54, 322-331.
92. Zheng, T., Zhu, Z., Wang, Z., Homer, R.J., Ma, B., Riese, R.J. Jr., Chapman, H.A. Jr., Shapiro, S.D., and Elias, J.A. (2000). Inducible targeting of IL-13 to the adult lung causes matrix metalloproteinase- and cathepsin-dependent emphysema. J. Clin. Invest. 106, 1081-1093.