Control of type IV collagenase activity by the urokinase-plasmin system: A regulatory mechanism with cell-bound reactants

EMBO Journal

Volumn 16, Issue 9, 1997, Pages 2319-2332

  Mazzieri, Roberta ^a   Masiero, Laura ^b   Zanetta, Lucia ^a   Monea, Sara ^a   Onisto, Maurizio ^b   Garbisa, Spiridione ^b   Mignatti, Paolo ^a,c  

^a UNIVERSITY OF PAVIA   (Italy)

^b UNIVERSITY OF PADOVA   (Italy)

^c NEW YORK UNIVERSITY MEDICAL CENTER   (United States)

Author keywords

Activation; Cell surface; Gelatinase; Plasmin; Urokinase

Indexed keywords

CELL SURFACE PROTEIN; ENZYME PRECURSOR; GELATINASE; GELATINASE A; GELATINASE B; PLASMIN; PLASMINOGEN; PROTEINASE INHIBITOR; UROKINASE;

ANGIOGENESIS; ANIMAL CELL; ARTICLE; CANCER INVASION; CELL INVASION; CONTROLLED STUDY; DRUG EFFECT; ENZYME ACTIVATION; ENZYME ACTIVITY; ENZYME REGULATION; METASTASIS; MOUSE; NONHUMAN; PRIORITY JOURNAL; PROTEIN BINDING; PROTEIN DEGRADATION; PROTEIN SECRETION;

COLLAGENASES; ENZYME ACTIVATION; ENZYME PRECURSORS; GELATINASES; HUMANS; MATRIX METALLOPROTEINASE 2; MATRIX METALLOPROTEINASE 9; METALLOENDOPEPTIDASES; MOLECULAR WEIGHT; PLASMIN; PLASMINOGEN ACTIVATORS; RECEPTORS, CELL SURFACE; SURFACE PROPERTIES; TUMOR CELLS, CULTURED; URINARY PLASMINOGEN ACTIVATOR;

ANIMALIA;

EID: 0030977009     PISSN: 02614189     EISSN: None     Source Type: Journal    
DOI: 10.1093/emboj/16.9.2319     Document Type: Article

References (92)

1. Aimes, R. T. and Quigley, J.P. (1995) Matrix metalloproteinase-2 is an interstitial collagenase. J. Biol. Chem., 270, 5872-5876.
2. Appella, E., Robinson, E.A., Ulrich, S.J., Stoppelh, A., Corti, M.P., Cassani, G. and Blasi, F. (1987) The receptor-binding sequence of urokinase. A biological function for the growth factor module of proteases. J. Biol. Chem., 262, 4437-4440.
3. Atkinson, S.J., Ward, R.V., Reynolds, J.J. and Murphy, G. (1992) Cell-mediated degradation of type IV collagen and gelatin films is dependent on the activation of matrix metalloproteinases. Biochem. J., 288, 605-611.
4. Baricos, W.H., Cortez, S.L. el-Dahr, S.S. and Schnaper, H.W. (1995) ECM degradation by cultured human mesangial cells is mediated by a PA/plasmin/MMP-2 cascade. Kidney Int. 47, 1039-47.
5. 2-terminal sequence and glycosylation variants. J. Biol. Chem., 265, 6453-6460.
6. r 92,00 gelatinase correlates with the metastatic phenotype in transformed rat embryo cells. Cancer Res., 50, 3872-3877.
7. Bernhard, E.J., Gruber, S.B. and Muschel, R.J. (1994) Direct evidence linking expression of matrix metalloproteinase 9 (92-kD gelatinase/. collagenase) to the metastatic phenotype in transformed rat embryo cells. Proc. Natl Acad. Sci. USA. 91, 4293-4297.
8. Brooks, P.C., Stromblad, S., Sanders, L.C., von Schalscha, T.L., Aimes, R.T., Stetler-Stevenson, W.G., Quigley, J.P. and Cheresh, D.A. (1996) Localization of matrix metalloproteinase MMP-2 to the surface of invasive cells by interaction with integrin alpha v beta 3. Cell. 85, 683-693.
9. r 72,000 type IV collagenase and interstitial collagenase in human tumorigenic cell lines. Cancer Res., 50, 6184-6191.
10. Cajot, J.-F., Schleuning, W.-D., Medcalf, R.L., Bamat, J., Testuz, J., Liebermann, L. and Sordat, B. (1989) Mouse L cells expressing human prourokinase-type plasminogen activator: effects on extracellular matrix desradation and invasion. J. Cell Biol., 109, 915-925.
11. Cajot, J.-F., Bamat, J., Bergonzelli, G.E., Kruithof, E.K.O., Medcalt, R.L., Testuz, J. and Sordat, B. (1990) Plasminogen-activator inhibitor type 1 is a potent natural inhibitor of extracellular matrix degradation by fibrosarcoma and colon carcinoma cells. Proc. Natl Acad. Sci. USA, 87, 6939-6943.
12. Cao, J., Sato, H., Takino, T. and Seiki, M. (1995) The C-terminal region of membrane type matrix metalloproteinase is a functional transmembrane domain required for pro-gelatinase A activation. J. Biol. Chem., 270, 801-805.
13. Collier, I.E. et al. (1988) H-ras oncogene-transformed human bronchial epithelial cells (TBE-1) secrete a single metalloproteinase capable of degrading basement membrane collagen. J. Biol. Chem., 263, 6579-6587.
14. Crawford, H.C. and Matrisian, L.M. (1995) Tumor and stromal expression of matrix metalloproteinases and their role in tumor progression. Invasion Metastasis, 14, 234-245.
15. Desrivieres, D., Lu, H., Peyri, N., Soria, C. Legrand, Y. and Menashi, S. (1993) Activation of the 92 kDa type IV collagenase by tissue kallikrein. J. Cell. Physiol., 157, 587-593.
16. Docherty, A.J.P., O'Connel, J., Grabble, T., Angal, S. and Murphy, G. (1992) The matrix metalloproteinases and their natural inhibitors: prospects for treating degenerative tissue diseases. Trends Biotechnol., 10, 200-207.
17. Ellis, V., Behrendt, N. and Danø, K. (1991) Plasminogen activation by receptor-bound urokinase. A kinetic study with both cell-associated and isolated receptor. J. Biol. Chem., 266, 12752-12758.
18. Emonard, H.P., Remacle, A.G., Noël, A.C. Grimaud, J.A., Stetler-Stevenson, W.G. and Foidart, J.M. (1992) Tumor cell surface associated binding site for M(r) 72,000 type IV collagenase. Cancer Res., 52, 5845-5848.
19. Felez, J., Chanquia, C.J., Fabregas, P., Plow, E.F. and Miles, L.A. (1993) Competition between plasminogen and tissue plasminogen activator for cellular binding sites. Blood, 82, 2433-2441.
20. Fridman, R. et al. (1993) Expression of human recombinant 72 kDa gelatinase and tissue inhibitor of metalloproteinase-2 (TIMP-2): characterization of complex and free enzyme. Biochem. J., 289, 411-416.
21. Garbisa, S., Kniska, K., Tryggvason, K., Foltz, C.M. and Liotta, L.A. (1980) Quantitation of basement membrane collagen degradation by living tumor cells in vitro. Cancer Letters, 9, 359-366.
22. Garbisa, S., Pozzatti, R., Mushel, R., Saffiotti, U., Ballin, M., Goldfarb, R., Khoury, G. and Liotta, L.A. (1987) Secretion of type IV collagenolytic protease and metastatic phenotype: induction by transfection with c-Ha-ras but not c-Ha-ras + Ad2-Ela. Cancer Res., 47, 1523-1528.
23. Garbisa, S., Scagliotti, G., Masiero, L., Di Francesco, C., Caenazzo, C., Onisto, M., Micela, M., Stetler-Stevenson, W.G. and Liotta, L.A. (1992) Correlation of serum metalloproteinase levels with lung cancer metastasis and response to therapy. Cancer Res., 52, 4548-4549.
24. Goldberg, G.I., Marmer, B.L., Grant, G.A., Eisen, A.Z., Wilhelm, A. and He, C. (1989) Human 72-kilodalton type IV collagenase forms a complex with a tissue inhibitor of metalloproteinases designated TIMP-2. Proc. Natl Acad. Sci. USA, 86, 8207-8211.
25. Goldberg, G.I., Strongin, A., Collier, I.E., Genrich, L.T. and Marmer, B.L. (1992) Interaction of 92-kD type IV collagenase with the tissue inhibitor of metalloproteinases prevents dimenzation, complex formation with interstitial collagenase, and activation of the proenzyme with stromelysin. J. Biol. Chem., 267, 4583-4591.
26. Gross, J.L., Moscatelli, D., Jaffe, E.A. and Rifkin, D.B. (1982) Plasminogen activator and collagenase production by cultured capillary endothelial cells. J. Cell Biol., 95, 974-981.
27. Hajjar, K.A. (1991) The endothelial cell tissue plasminogen activator receptor. Specific interaction with plasminogen. J. Biol. Chem., 266, 21962-21970.
28. Hajjar, K.A., Harpel, P.C., Jaffe, E.A. and Nachman, R.L. (1986) Binding of plasminogen to cultured human endothelial cells. J. Biol. Chem., 261, 11656-11662.
29. Hajjar, K.A., Jacovina, A.T. and Chacko, J. (1994) An endothelial cell receptor for plasminogen/tissue plasminogen activator. I. Identity with annexin II. J. Biol. Chem., 269, 21191-21197.
30. He, C. Wilhelm, S.M., Pentland, A.P., Marmer, B.L., Grant, G.A., Eisen, A.Z. and Goldberg, G.I. (1989) Tissue cooperation in a proteolytic cascade activating human interstitial collagenase. Proc. Natl Acad. Sci. USA, 86, 2632-2636.
31. Hébert, C.A. and Baker, J.B. (1988) Linkage of extracellular plasminogen activator to the fibroblast cytoskeleton: colocalization of cell surface urokinase with vinculin. J. Cell Biol., 106, 1241-1248.
32. Heussen, C. and Dowdle, E.B. (1980) Electrophoretic analysis of plasminogen activators in polyacrylamide gels containing sodium dodecyl sulfate and copolymerized substrates. Anal. Biochem., 102, 196-202.
33. Himelstein, B.P., Canete-Soler, R., Bernhard, E.J., Dilks, D.W. and Muschel, R.L. (1995) Metalloproteinases in tumor progression: the contribution of MMP-9. Invest. Methods, 14, 246-258.
34. Isseroff, R.R. and Rifkin, D.B. (1983) Plasminogen is present in the basal layer of epidermis. J. Invest. Dermatol., 80, 297-299.
35. Itoh, Y., Binner, S. and Nagase, H. (1995) Steps involved in activation of the complex of pro-matrix metalloproteinase 2 (progelatinase A) and tissue inhibitor of metalloproteinases (TIMP)-2 by 4-aminophenylmercuric acetate. Biochem. J., 308, 645-651.
36. Juarez, J., Clayman, G., Nakajima, M., Tanabe, K.K., Saya, H., Nicolson, G.L. and Boyd, D. (1993) Role and regulation of expression of 92-kD type IV collagenase (MMP-9) in 2 invasive squamous-cell-carcinoma cell lines of the oral cavity. Int. J. Cancer, 55, 10-18.
37. Kanalas, J.J. and Makker, S.P. (1991) Identification of the rat Heymann nephritis autoantigen (gp330) as a receptor site for plasminogen. J. Biol. Chem., 266, 10825-10829.
38. Keski-Oja, J., Lohi, J., Tuuttila, A., Tryggvason, K. and Vartio, T. (1992) Proteolytic processing of the 72,000-Da type IV collagenase by urokinase plasminogen activator. Exp. Cell Res., 202, 471-476.
39. Kleiner, D.E. and Stetler-Stevenson, W.G. (1993) Structural biochemistry and activation of matrix metalloproteinases. Curr. Opin. Cell Biol., 5, 891-897.
40. Knudsen, B.S., Silverstein, R.L., Leung, L.L.K., Harpel, P.C. and Nachman, R.L. (1986) Binding of plasminogen to extracellular matrix. J. Biol. Chem., 261, 10765-10771.
41. Kohn, E.C. and Liotta, L.A. (1995) Molecular insights into cancer invasion: strategies for prevention and intervention. Cancer Res., 55, 1856-1862.
42. Levy, A.T., Cioce, V., Sobel, M.E., Garbisa, S., Grigioni, W., Liotta, L.A. and Stetler-Stevenson, W.G. (1991) Increased expression of the 72 kDa type IV collagenase in human colonie adenocarcinoma. Cancer Res., 51, 439-444.
43. Librach, C.L. et al. (1991) 92-kD type IV collagenase mediates invasion of human cytotrophoblasts. J. Cell Biol., 113, 437-149.
44. Lohi, J. and Keski-Oja, J. (1995) Calcium ionophores decrease pericellular gelatinolytic activity via inhibition of 92-kDa gelatinase expression and decrease of 72-kDA gelatinase activation. J. Biol. Chem., 270, 17602-17609.
45. Matrisian, L.M. (1990) Metalloproteinases and their inhibitors in matrix remodeling. Trends Genet., 6, 121-125.
46. Mazzieri, R., Høyer-Hansen, G., Rønne, E., Leber, D., Vagnarelli, P., Raimondi, E., De Carli, L., Dano, K. and Mignatti, P. (1994) Urokinase and urokinase receptor expression in somatic cell hybrids. Fibrinolysis, 8, 344-352.
47. Mignatti, P. and Rifkin, D.B. (1993) Biology and biochemistry of proteinases in tumor invasion. Physiol. Rev., 73, 161-195.
48. Mignatti, P. Robbins, E. and Rifkin, D.B. (1986) Tumor invasion through the human amniotic membrane: requirement for a proteinase cascade. Cell, 47, 487-498.
49. Mignatti, P., Tsuboi, R., Robbins, E. and Rifkin, D.B. (1989) In vitro angiogenesis on the human amniotic membrane: requirement for basic fibroblast growth factor-induced proteinases. J. Cell Biol., 108, 671-682.
50. Mignatti, P. Mazzieri, R. and Rifkin, D.B. (1991) Expression of the urokinase receptor in vascular endothelial cells is stimulated by basic fibroblast growth factor. J. Cell Biol., 113, 1193-1201.
51. Miles, L.A. and Plow, E.F. (1985) Binding and activation of plasminogen on the platelet surface. J. Biol. Chem., 260, 4303-4311.
52. Monsky, W.L., Kelly, T., Lin, C.Y., Yeh, Y., Stetler-Stevenson, W.G., Mueller, S.C. and Chen, W.T. (1993) Binding and localization of M(r) 72,000 matrix metalloproteinase at cell surface invadopodia. Cancer Res., 53, 3159-64.
53. Montgomery, A.M., De Clerck, Y.A., Langley, K.E., Reisfeld, B.M. and Mueller, B.M. (1993) Melanoma-mediated dissolution of extracellular matrix: contribution of urokinase-dependent and metalloproteinase-dependent proteolytic pathways. Cancer Res., 53, 693-700.
54. Moscatelli, D. and Rifkin, D.B. (1988) Membrane and matrix localization of proteinases: a common theme in tumor cell invasion and angiogenesis. Biochim. Biophys. Acta, 948, 67-85.
55. Murphy, G., Atkinson, S., Ward, R., Gavrilovic, J. and Reynolds, J.J. (1992a) The role of plasminogen activators in the regulation of connective tissue metalloproteinases. Ann. NY Acad. Sci., 667, 1-12.
56. Murphy, G., Willenbrock, F., Ward, R.V., Cockett, M.I., Eaton, D. and Docherty, A.J. (1992b) The C-terminal domain of 72 kDa gelatinase A is not required for catalysis, but is essential for membrane activation and modulates interactions with tissue inhibitors of metalloproteinases. Biochem. J., 283, 637-641.
57. Nagase, H., Ogata, Y., Suzuki, K., Enghild, J.J. and Salvensen, G. (1991) Substrate specificities and activation mechanisms of matrix metalloproteinases. Biochem. Soc. Trans., 19, 715-718.
58. Nagase, H., Suzuki, K., Morodomi, T., Enghild, J.J. and Salvesen, G. (1992) Activation mechanisms of the precursor of matrix metalloproteinases 1, 2 and 3. Matrix Suppl., 1, 237-244.
59. Nomura, H., Sato, H., Seiki, M., Mai, M., and Okada, Y. (1995) Expression of membrane-type matrix metalloproteinase in human gastric carcinomas. Cancer Res., 55, 3263-3266.
60. Okada, Y., Morodomi, T., Enghild, J.J., Suzuki, K., Yasui, A., Nakanishi, I., Salvesen, G. and Nagase, H. (1990) Matrix metalloproteinase 2 from human rheumatoid synovial fibroblasts. Purification and activation of the precursor and enzymic properties. Eur. J. Biochem., 194, 721-730.
61. Okada, Y., Gonoji, Y., Naka, K., Tomita, K., Nakanishi, I., Iwata, K., Yamashita, K. and Hayakawa, T. (1992) Matrix metalloproteinase 9 (92-kD gelatinase/type IV collagenase) from HT1080 human fibrosarcoma cells. Purification and activation of the precursor and enzymic properties. J. Biol. Chem., 267, 21712-21719.
62. Okada, Y., Bellocq, J.R. Rouyer, N., Chenard, M.P, Rio, M.C., Chambon, P. and Basset, P. (1995) Membrane-type matrix metalloproteinase (MT-MMP) gene is expressed in stromal cells of human colon, breast, head and neck carcinomas. Proc. Natl Acad. Sci. USA, 92, 2730-2734.
63. Onisto, M., Garbisa, S., Caenazzo, C., Freda, M., Di Francesco, C., Liotta, L.A. and Stetler-Stevenson, W.G. (1993) RT-PCR phenotyping of metalloproteinases and inhibitors involved in tumor matrix invasion. Diagn. Mol. Pathol., 2, 74-80.
64. Pedersen, N., Masucci, M.T., Moller, L.B. and Blasi, F. (1991) A non-catalytic human urokinase plasminogen activator derivative produced by mouse cells has full receptor-binding activity. Fibrinolysis, 5, 155-164.
65. Petersen, L.C., Lund, L.R., Nielsen, L.S., Dana, K. and Skriver, L. (1988) One chain urokinase-type plasminogen activator from human sarcoma cells is a proenzyme with little or no intrinsic activity. J. Biol. Chem., 263, 11189-11195.
66. Plow, E.F. and Miles, L.A. (1990) Plasminogen receptors in the mediation of pericellular proteolysis. Cell. Differ. Dev., 32, 293-298.
67. Plow, E.F., Freaney, D.E., Plescia, J. and Miles, L.A. (1986) The plasminogen system and cell surfaces: evidence for plasminogen and urokinase receptors on the same cell type. J. Cell. Biol., 103, 2411-2420.
68. Plow, E.F., Herren, T., Redlitz, A., Miles, L.A. and Hoover-Plow, J.L. (1995) The cell biology of the plasminogen system. FASEB J., 9, 939-945.
69. Pöllanen, J., Hedman, K., Nielsen, L.S., Danø, K. and Vaheri, A. (1988) Ultrastructural localization of plasma membrane-associated urokinase-type plasminogen activator at focal contacts. J. Cell Biol., 106, 87-95.
70. Reith, A. and Rucklidge, G.J. (1992) Invasion of brain tissue by primary glioma: evidence for the involvement of urokinase-type plasminogen activator as an activator of type IV collagenase. Biochem. Biophys. Res. Commun., 186, 348-354.
71. Robbins, K.C. and Summaria, L. (1970) Human plasminogen and plasmin. Methods Enzymol., 19, 184-199.
72. Roldan, A.L., Cubellis, M.V. Masucci, M.T., Behrendt, N., Lund, L.R., Danø, K., Appella, E. and Blasi, F. (1990) Cloning and expression of the receptor for human urokinase plasminogen activator, a central molecule in cell surface, plasmin dependent proteolysis. EMBO J., 9, 467-474.
73. Saksela, O. and Rifkin, D.B. (1988) Cell-associated plasminogen activation: regulation and physiological functions. Annu. Rev. Cell Biol., 4, 93-126.
74. Sato, H., Takino, T., Okada, Y., Cao, J., Shinagawa, A., Yamamoto, E. and Seiki, M. (1994) A matrix metalloproteinase expressed on the surface of invasive tumor cells. Nature, 370, 61-65.
75. Shapiro, S.D., Griffin, G.L., Gilbert, D.J., Jenkins, N.A., Copeland, N.G., Welgus, H.G., Senior, R.M. and Ley, T.J. (1992) Molecular cloning, chromosomal localization, and bacterial expression of a Murine Macrophage Metalloelastase. J. Biol. Chem., 267, 4664-4671.
76. Stetler-Stevenson, W.G. (1995) Progelatinase A activation during tumor cell invasion. Invest. Methods, 14, 259-268.
77. Stetler-Stevenson, W.G., Krutzsch, H.C., Wacher, M.P. Margulies, I.M.K. and Liotta, L.A. (1989) The activation of human type IV collagenase proenzyme. Sequence identification of the major conversion product following organomercurial activation. J. Biol. Chem., 264, 1353-1356.
78. Stetler-Stevenson, W.G., Aznavoorian, S. and Liotta, L.A. (1993) Tumor cell interactions with the extracellular matrix during invasion and metastasis. Annu. Rev. Cell Biol., 9, 541-573.
79. Stoppelli, M.P., Corti, A., Soffientini, A., Cassani, G., Blasi, F. and Assoian, R.K. (1985) Differentiation-enhanced binding of the aminoterminal fragment of human urokinase plasminogen activator to a specific receptor on U937 monocytes. Proc. Natl Acad. Sci. USA. 82, 4939-4943.
80. Stoppelli, M.P., Tacchetti, C., Cubellis, M.V., Corti, A., Hearing, V.J., Cassani, G., Appella, E. and Blasi, F. (1986) Autocrine saturation of pro-urokinase receptors on human A431 cells. Cell, 45, 675-684.
81. Strongin, A.Y. Marmer, B.L., Grant, G.A. and Goldberg, G.I. (1993) Plasma membrane-dependent activation of the 72-kD type IV collagenase is prevented by complex formation with TIMP-2. J. Biol. Chem., 268, 14033-14039.
82. Strongin, A.Y, Collier, I., Bannikov, G., Marmer, B.N., Grant, G.A. and Goldberg, G.I. (1995) Mechanism of cell surface activation of 72-kDa type IV collagenase. Isolation of the activated form of the membrane metalloproteinase. J. Biol. Chem., 270, 5331-5338.
83. Takino, T., Sato, H., Shinagawa, A. and Seiki, M. (1995) Identification of the second membrane-type metalloproteinase (MT-MMP-2) gene from a human placenta cDNA library. J. Biol. Chem., 270, 23013-23020.
84. Unkeless, J.C., Tobia, A., Ossowski, L., Quigley, P., Rifkin, D.B. and Reich, E. (1973) An enzymatic function associated with transformation of fibroblasts with oncogenic viruses. I. Chick embryo fibroblast cultures transformed by avian RNA tumor viruses. J. Exp. Med., 137, 85-111.
85. r 55,000 form of the human plasminogen activator, urokinase. J. Cell Biol., 100, 86-92.
86. Wacher, M.P., Krutzsch, H.C., Liotta, L.A. and Stetler-Stevenson, W.G. (1990) Development of a novel substrate capture immunoassay for the detection of a neutral metalloproteinase capable of degrading basement membrane (type IV) collagen. J. Immunol. Methods, 126, 239-245.
87. Ward, R.V., Atkinson, S.J., Slocombe, P.M., Docherty, A.J.R. Reynolds, J.J. and Murphy, G. (1991) Tissue inhibitor of metalloproteinases-2 inhibits the activation of 72 kDa progelatinase by fibroblast membranes. Biochim. Biophys. Acta, 1079, 242-246.
88. Watanabe, H., Nakanishi, L. Yamashita, K., Hayakawa, T. and Okada, Y. (1993) Matrix metalloproteinase-9 (92 kDa gelatinase/type IV collagenase) from U937 monoblastoid cells: correlation with cellular invasion. J. Cell Sci., 104, 991-999.
89. Welgus, H.G., Jeffrey, J.J. and Eisen, A.Z. (1981) The collagen substrate specificity of human skin fibroblast collagenase. J. Biol. Chem., 256, 9511-9515.
90. Werb, Z., Mainardi, C., Vater, C. and Harris, E.D. (1977) Endogenous activation of latent collagenase by rheumatoid synovial cells. Evidence for a role of plasminogen activator. N. Engl. J. Med., 296, 1017-1023.
91. Wilhelm, S.M., Collier, I.E., Marmer, B.L., Eisen, A.Z., Grant, G.A. and Goldberg, G.I. (1989) SV40-transformed human lung fibroblasts secrete a 92-kTJ type IV collagenase which is identical to that secreted by normal human macrophages. J. Biol. Chem., 264, 17213-17221.
92. Wong, A.R, Cortez, S.L. and Baricos, W.H. (1992) Role of plasmin and gelatinase in extracellular matrix degradation by cultured rat mesangial cells. Am. J. Physiol. 263, 1112-1118.