Kallikrein-related peptidases and inhibitors of the skin

Characterization, regulation, and interactions within the protease web

Volumn , Issue , 2012, Pages 329-348

  Brattsand, Maria ^a  

^a UMEÅ UNIVERSITY   (Sweden)

Author keywords

[No Author keywords available]

Indexed keywords

EID: 84976223101     PISSN: None     EISSN: None     Source Type: Book    
DOI: None     Document Type: Chapter

References (103)

1. Ahmed, A., Kandola, P., Ziada, G., and Parenteau, N. (2001). Purification and partial amino acid sequence of proteins from human epidermal keratinocyte conditioned medium. J. Protein Chem. 20, 273-278
2. Bitoun, E., Micheloni, A., Lamant, L., Bonnart, C., Tartaglia-Polcini, A., Cobbold, C., Al Saati, T., Mariotti, F., Mazereeuw-Hautier, J., Boralevi, F., Hohl, D., Harper, J., Bodemer, C., D'Alessio, M., and Hovnanian, A. (2003). LEKTI proteolytic processing in human primary keratinocytes, tissue distribution and defective expression in Netherton syndrome. Hum. Mol. Genet. 12, 2417-2430
3. Blaber, S.I., Yoon, H., Scarisbrick, I.A., Juliano, M.A., and Blaber, M. (2007). The autolytic regulation of human kallikrein-related peptidase 6. Biochemistry 46, 5209-5217
4. Borgoño, C.A., Michael, I.P., Komatsu, N., Jayakumar, A., Kapadia, R., Clayman, G.L., Sotiropoulou, G., and Diamandis, E.P. (2007). A potential role for multiple tissue kallikrein serine proteases in epidermal desquamation. J. Biol. Chem. 282, 3640-3652
5. Bowcock, A.M., Shannon, W., Du, F., Duncan, J., Cao, K., Aftergut, K., Catier, J., Fernandez-Vina, M.A., and Menter, A. (2001). Insights into psoriasis and other inflammatory diseases from large-scale gene expression studies. Hum. Mol. Genet. 10, 1793-1805
6. Brattsand, M., and Egelrud, T. (1999). Purification, molecular cloning, and expression of a human stratum corneum trypsin-like serine protease with possible function in desquamation. J. Biol. Chem. 274, 30033-30040
7. Brattsand, M., Stefansson, K., Lundh, C., Haasum, Y., and Egelrud, T. (2005). A proteolytic cascade of kallikreins in the stratum corneum. J. Invest. Dermatol. 124, 198-203
8. Brattsand, M., Stefansson, K., Hubiche, T., Nilsson, S.K., and Egelrud, T. (2009). SPINK9: a selective, skin-specific Kazal-type serine protease inhibitor. J. Invest. Dermatol. 129, 1656-1665
9. Briot, A., Deraison, C., Lacroix, M., Bonnart, C., Robin, A., Besson, C., Dubus, P., and Hovnanian, A. (2009). Kallikrein 5 induces atopic dermatitis-like lesions through PAR2-mediated thymic stromal lymphopoietin expression in Netherton syndrome. J. Exp. Med. 206, 1135-1147
10. Brody, I. (1968). An electron-microscopic study of the junctional and regular desmosomes in normal human epidermia. Acta Derm. Venereol. 48, 290-302
11. Burns, T., Breathnach, S., Cox, N., and Griffiths, C., eds. (2004). Rook's Textbook of Dermatology (Blackwell Publishing)
12. Candi, E., Schmidt, R., and Melino, G. (2005). The cornified envelope: a model of cell death in the skin. Nat. Rev. Mol. Cell Biol. 6, 328-340
13. Caubet, C., Jonca, N., Brattsand, M., Guerrin, M., Bernard, D., Schmidt, R., Egelrud, T., Simon, M., and Serre, G. (2004). Degradation of corneodesmosome proteins by two serine proteases of the kallikrein family, SCTE/KLK5/hK5 and SCCE/KLK7/hK7. J. Invest. Dermatol. 122, 1235-1244
14. Chavanas, S., Bodemer, C., Rochat, A., Hamel-Teillac, D., Ali, M., Irvine, A., Bonafé, J., Wilkinson, J., Taïeb, A., Barrandon, Y., Harper, J.I., de Prost, Y., and Hovnanian, A. (2000). Mutations in SPINK5, encoding a serine protease inhibitor, cause Netherton syndrome. Nat. Genet. 25, 141-142
15. Comel, M. (1949). Ichtyosis linearis circumflexa. Dermatologica 98, 133-136
16. Cork, M.J., Danby, S.G., Vasilopoulos, Y., Hadgraft, J., Lane, M.E., Moustafa, M., Guy, R.H., Macgowan, A.L., Tazi-Ahnini, R., and Ward, S.J. (2009). Epidermal barrier dysfunction in atopic dermatitis. J. Invest. Dermatol. 129, 1892-1908
17. Deraison, C., Bonnart, C., Lopez, F., Besson, C., Robinson, R., Jayakumar, A., Wagberg, F., Brattsand, M., Hachem, J.P., Leonardsson, G., and Hovnanian, A. (2007). LEKTI fragments specifically inhibit KLK5, KLK7, and KLK14 and control desquamation through a pH-dependent interaction. Mol. Biol. Cell 18, 3607-3619
18. Descargues, P., Deraison, C., Bonnart, C., Kreft, M., Kishibe, M., Ishida-Yamamoto, A., Elias, P., Barrandon, Y., Zambruno, G., Sonnenberg, A., and Hovnanian, A. (2005). Spink5-deficient mice mimic Netherton syndrome through degradation of desmoglein 1 by epidermal protease hyperactivity. Nat. Genet. 37, 56-65
19. Descargues, P., Deraison, C., Prost, C., Fraitag, S., Mazereeuw-Hautier, J., D'Alessio, M., Ishida-Yamamoto, A., Bodemer, C., Zambruno, G., and Hovnanian, A. (2006). Corneodesmosomal cadherins are preferential targets of stratum corneum trypsin-and chymotrypsin-like hyperactivity in Netherton syndrome. J. Invest. Dermatol. 126, 1622-1632
20. Egelrud, T., Hofer, P.A., and Lundstrom, A. (1988). Proteolytic degradation of desmosomes in plantar stratum corneum leads to cell dissociation in vitro. Acta Derm. Venereol. 68, 93-97
21. Egelrud, T., and Lundstrom, A. (1991). A chymotrypsin-like proteinase that may be involved in desquamation in plantar stratum corneum. Arch Dermatol. Res. 283, 108-112
22. Egelrud, T. (2000). Desquamation in the stratum corneum. Acta Derm. Venereol. Suppl. (Stockh) 208, 44-45
23. Egelrud, T., Brattsand, M., Kreutzmann, P., Walden, M., Vitzithum, K., Marx, U.C., Forssmann, W.G., and Magert, H.J. (2005). hK5 and hK7, two serine proteinases abundant in human skin, are inhibited by LEKTI domain 6. Br. J. Dermatol. 153, 1200-1203
24. Eissa, A., Amodeo, V., Smith, C.R., and Diamandis, E.P. (2011). Kallikrein-related peptidase-8 (KLK8) is an active serine protease in human epidermis and sweat and is involved in a skin barrier proteolytic cascade. J. Biol. Chem. 286, 687-706
25. Ekholm, E., and Egelrud, T. (1998). The expression of stratum corneum chymotryptic enzyme in human anagen hair follicles: further evidence for its involvement in desquamation-like processes. Br. J. Dermatol. 139, 585-590
26. Ekholm, E., Sondell, B., Stranden, P., Brattsand, M., and Egelrud, T. (1998). Expression of stratum corneum chymotryptic enzyme in human sebaceous follicles. Acta Derm. Venereol. 78, 343-347
27. Ekholm, E., and Egelrud, T. (1999). Stratum corneum chymotryptic enzyme in psoriasis. Arch Dermatol. Res. 291, 195-200
28. Ekholm, E., Brattsand, M., and Egelrud, T. (1999). Stratum corneum tryptic enzyme-A missing link in desquamation? J. Invest. Dermatol. 113, 454-454
29. Ekholm, I.E., Brattsand, M., and Egelrud, T. (2000). Stratum corneum tryptic enzyme in normal epidermis: a missing link in the desquamation process? J. Invest. Dermatol. 114, 56-63
30. Elias, P.M. (2004). The epidermal permeability barrier: from the early days at Harvard to emerging concepts. J. Invest. Dermatol. 122, xxxvi-xxxix
31. Fortugno, P., Bresciani, A., Paolini, C., Pazzagli, C., El Hachem, M., D'Alessio, M., and Zambruno, G. (2011). Proteolytic activation cascade of the Netherton syndrome-defective protein, LEKTI, in the epidermis: implications for skin homeostasis. J. Invest. Dermatol. 131, 2223-2232
32. Franzke, C., Baici, A., Bartels, J., Christophers, E., and Wiedow, O. (1996). Antileukoprotease inhibits stratum corneum chymotryptic enzyme. Evidence for a regulative function in desquamation. J. Biol. Chem. 271, 21886-21890
33. Goettig, P., Magdolen, V., and Brandstetter, H. (2010). Natural and synthetic inhibitors of kallikreinrelated peptidases (KLKs). Biochimie 92, 1546-1567
34. Hachem, J.P., Crumrine, D., Fluhr, J., Brown, B.E., Feingold, K.R., and Elias, P.M. (2003). pH directly regulates epidermal permeability barrier homeostasis, and stratum corneum integrity/ cohesion. J. Invest. Dermatol. 121, 345-353
35. Hachem, J.P., Houben, E., Crumrine, D., Man, M.Q., Schurer, N., Roelandt, T., Choi, E.H., Uchida, Y., Brown, B.E., Feingold, K.R., and Elias, P.M. (2006). Serine protease signaling of epidermal permeability barrier homeostasis. J. Invest. Dermatol. 126, 2074-2086
36. Haftek, M., Simon, M., Kanitakis, J., Marechal, S., Claudy, A., Serre, G., and Schmitt, D. (1997). Expression of corneodesmosin in the granular layer and stratum corneum of normal and diseased epidermis. Br. J. Dermatol. 137, 864-873
37. Hanson, K.M., Behne, M.J., Barry, N.P., Mauro, T.M., Gratton, E., and Clegg, R.M. (2002). Two-photon fluorescence lifetime imaging of the skin stratum corneum pH gradient. Biophys. J. 83, 1682-1690
38. Hansson, L., Stromqvist, M., Backman, A., Wallbrandt, P., Carlstein, A., and Egelrud, T. (1994). Cloning, expression, and characterization of stratum corneum chymotryptic enzyme. A skin-specific human serine proteinase. J. Biol. Chem. 269, 19420-19426
39. Hansson, L., Backman, A., Ny, A., Edlund, M., Ekholm, E., Ekstrand Hammarstrom, B., Tornell, J., Wallbrandt, P., Wennbo, H., and Egelrud, T. (2002). Epidermal overexpression of stratum corneum chymotryptic enzyme in mice: a model for chronic itchy dermatitis. J. Invest. Dermatol. 118, 444-449
40. Hubiche, T., Ged, C., Benard, A., Leaute-Labreze, C., McElreavey, K., de Verneuil, H., Taieb, A., and Boralevi, F. (2007). Analysis of SPINK 5, KLK 7 and FLG genotypes in a French atopic dermatitis cohort. Acta Derm. Venereol. 87, 499-505
41. Ishida-Yamamoto, A., Simon, M., Kishibe, M., Miyauchi, Y., Takahashi, H., Yoshida, S., O'Brien, T.J., Serre, G., and Iizuka, H. (2004). Epidermal lamellar granules transport different cargoes as distinct aggregates. J. Invest. Dermatol. 122, 1137-1144
42. Ishida-Yamamoto, A., Deraison, C., Bonnart, C., Bitoun, E., Robinson, R., O'Brien, T.J., Wakamatsu, K., Ohtsubo, S., Takahashi, H., Hashimoto, Y., Dopping-Hepenstal, P.J., McGrath, J.A., Iizuka, H., Richard, G., and Hovnanian, A. (2005). LEKTI is localized in lamellar granules, separated from KLK5 and KLK7, and is secreted in the extracellular spaces of the superficial stratum granulosum. J. Invest. Dermatol. 124, 360-366
43. Jayakumar, A., Kang, Y., Henderson, Y., Mitsudo, K., Liu, X., Briggs, K., Wang, M., Frederick, M.J., El-Naggar, A.K., Bebök, Z., and Clayman, G.L. (2005). Consequences of C-terminal domains and N-terminal signal peptide deletions on LEKTI secretion, stability, and subcellular distribution. Arch. Biochem. Biophys. 435, 89-102
44. Kanitakis, J., Lora, V., Chouvet, B., Zambruno, G., Haftek, M., and Faure, M. (2011). Circumscribed palmo-plantar hypokeratosis: a disease of desquamation? Immunohistological study of five cases and literature review. J. Eur. Acad. Dermatol. Venereol. 25, 296-301
45. Kirihara, T., Matsumoto-Miyai, K., Nakamura, Y., Sadayama, T., Yoshida, S., and Shiosaka, S. (2003). Prolonged recovery of ultraviolet B-irradiated skin in neuropsin (KLK8)-deficient mice. Br. J. Dermatol. 149, 700-706
46. Kishibe, M., Bando, Y., Terayama, R., Namikawa, K., Takahashi, H., Hashimoto, Y., Ishida-Yamamoto, A., Jiang, Y.P., Mitrovic, B., Perez, D., Iizuka, H., and Yoshida, S. (2007). Kallikrein 8 is involved in skin desquamation in cooperation with other kallikreins. J. Biol. Chem. 282, 5834-5841
47. Komatsu, N., Takata, M., Otsuki, N., Ohka, R., Amano, O., Takehara, K., and Saijoh, K. (2002). Elevated stratum corneum hydrolytic activity in Netherton syndrome suggests an inhibitory regulation of desquamation by SPINK5-derived peptides. J. Invest. Dermatol. 118, 436-443
48. Komatsu, N., Takata, M., Otsuki, N., Toyama, T., Ohka, R., Takehara, K., and Saijoh, K. (2003). Expression and localization of tissue kallikrein mRNAs in human epidermis and appendages. J. Invest. Dermatol. 121, 542-549
49. Komatsu, N., Saijoh, K., Sidiropoulos, M., Tsai, B., Levesque, M., Elliott, M., Takehara, K., and Diamandis, E. (2005a). Quantification of human tissue kallikreins in the stratum corneum: dependence on age and gender. J. Invest. Dermatol. 125, 1182-1189
50. Komatsu, N., Saijoh, K., Toyama, T., Ohka, R., Otsuki, N., Hussack, G., Takehara, K., and Diamandis, E. (2005b). Multiple tissue kallikrein mRNA and protein expression in normal skin and skin diseases. Br. J. Dermatol. 153, 274-281
51. Komatsu, N., Suga, Y., Saijoh, K., Liu, A.C., Khan, S., Mizuno, Y., Ikeda, S., Wu, H.K., Jayakumar, A., Clayman, G.L., Shirasaki, F., Takehara, K., and Diamandis, E.P. (2006). Elevated human tissue kallikrein levels in the stratum corneum and serum of peeling skin syndrome-type B patients suggests an over-desquamation of corneocytes. J. Invest. Dermatol. 126, 2338-2342
52. Komatsu, N., Saijoh, K., Kuk, C., Liu, A.C., Khan, S., Shirasaki, F., Takehara, K., and Diamandis, E.P. (2007a). Human tissue kallikrein expression in the stratum corneum and serum of atopic dermatitis patients. Exp. Dermatol. 16, 513-519
53. Komatsu, N., Saijoh, K., Kuk, C., Shirasaki, F., Takehara, K., and Diamandis, E.P. (2007b). Aberrant human tissue kallikrein levels in the stratum corneum and serum of patients with psoriasis: dependence on phenotype, severity and therapy. Br. J. Dermatol. 156, 875-883
54. Komatsu, N., Saijoh, K., Jayakumar, A., Clayman, G.L., Tohyama, M., Suga, Y., Mizuno, Y., Tsukamoto, K., Taniuchi, K., Takehara, K., and Diamandis, E.P. (2008). Correlation between SPINK5 gene mutations and clinical manifestations in Netherton syndrome patients. J. Invest. Dermatol. 128, 1148-1159
55. Kreutzmann, P., Schulz, A., Standker, L., Forssmann, W.G., and Mägert, H.J. (2004). Recombinant production, purification and biochemical characterization of domain 6 of LEKTI: a temporary Kazal-type-related serine proteinase inhibitor. J. Chromatogr. B Analyt. Technol. Biomed. Life Sci. 803, 75-81
56. Lai, Y., and Gallo, R.L. (2009). AMPed up immunity: how antimicrobial peptides have multiple roles in immune defense. Trends Immunol. 30, 131-141
57. Lundqvist, E.N., and Egelrud, T. (1997). Biologically active, alternatively processed interleukin-1 beta in psoriatic scales. Eur. J. Immunol. 27, 2165-2171
58. Lundstrom, A., and Egelrud, T. (1990). Evidence that cell shedding from plantar stratum corneum in vitro involves endogenous proteolysis of the desmosomal protein desmoglein I. J. Invest. Dermatol. 94, 216-220
59. Lundstrom, A., Serre, G., Haftek, M., and Egelrud, T. (1994). Evidence for a role of corneodesmosin, a protein which may serve to modify desmosomes during cornification, in stratum corneum cell cohesion and desquamation. Arch Dermatol. Res. 286, 369-375
60. Lundwall, A., and Clauss, A. (2002). Identification of a novel protease inhibitor gene that is highly expressed in the prostate. Biochem. Biophys. Res. Commun. 290, 452-456
61. Luo, L.Y., and Jiang, W. (2006). Inhibition profiles of human tissue kallikreins by serine protease inhibitors. Biol. Chem. 387, 813-816
62. Mägert, H.J., Standker, L., Kreutzmann, P., Zucht, H.D., Reinecke, M., Sommerhoff, C.P., Fritz, H., and Forssmann, W.G. (1999). LEKTI, a novel 15-domain type of human serine proteinase inhibitor. J. Biol. Chem. 274, 21499-21502
63. McNeely, T.B., Dealy, M., Dripps, D.J., Orenstein, J.M., Eisenberg, S.P., and Wahl, S.M. (1995). Secretory leukocyte protease inhibitor: a human saliva protein exhibiting anti-human immunodeficiency virus 1 activity in vitro. J. Clin. Invest. 96, 456-464
64. Meyer-Hoffert, U., Wu, Z., and Schröder, J.M. (2009). Identification of lympho-epithelial Kazal-type inhibitor 2 in human skin as a kallikrein-related peptidase 5-specific protease inhibitor. PLoS ONE 4, e4372
65. Meyer-Hoffert, U., Wu, Z., Kantyka, T., Fischer, J., Latendorf, T., Hansmann, B., Bartels, J., He, Y., Glaser, R., and Schröder, J.M. (2010). Isolation of SPINK6 in human skin: selective inhibitor of kallikrein-related peptidases. J. Biol. Chem. 285, 32174-32181
66. Netherton, E.W. (1958). A unique case of trichorrhexis nodosa; bamboo hairs. AMA Arch. Derm. 78, 483-487
67. Nylander-Lundqvist, E., and Egelrud, T. (1997). Formation of active IL-1 beta from pro-IL-1 beta catalyzed by stratum corneum chymotryptic enzyme in vitro. Acta Derm. Venereol. 77, 203-206
68. Ohler, A., Debela, M., Wagner, S., Magdolen, V., and Becker-Pauly, C. (2010). Analyzing the protease web in skin: meprin metalloproteases are activated specifically by KLK4, 5 and 8 vice versa leading to processing of proKLK7 thereby triggering its activation. Biol. Chem. 391, 455-460
69. Ohman, H., and Vahlquist, A. (1994). In vivo studies concerning a pH gradient in human stratum corneum and upper epidermis. Acta Derm. Venereol. 74, 375-379
70. Oikonomopoulou, K., Hansen, K.K., Saifeddine, M., Vergnolle, N., Tea, I., Blaber, M., Blaber, S.I., Scarisbrick, I., Diamandis, E.P., and Hollenberg, M.D. (2006). Kallikrein-mediated cell signalling: targeting proteinase-activated receptors (PARs). Biol. Chem. 387, 817-824
71. Pearton, D.J., Nirunsuksiri, W., Rehemtulla, A., Lewis, S.P., Presland, R.B., and Dale, B.A. (2001). Proprotein convertase expression and localization in epidermis: evidence for multiple roles and substrates. Exp. Dermatol. 10, 193-203
72. Raghunath, M., Tontsidou, L., Oji, V., Aufenvenne, K., Schurmeyer-Horst, F., Jayakumar, A., Stander, H., Smolle, J., Clayman, G.L., and Traupe, H. (2004). SPINK5 and Netherton syndrome: novel mutations, demonstration of missing LEKTI, and differential expression of transglutaminases. J. Invest. Dermatol. 123, 474-483
73. Raknerud, N. (1975). The ultrastructure of the interfollicular epidermis of the hairless (hr/hr) mouse. III. Desmosomal transformation during keratinization. J. Ultrastruct. Res. 52, 32-51
74. Roedl, D., Traidl-Hoffmann, C., Ring, J., Behrendt, H., and Braun-Falco, M. (2009). Serine protease inhibitor lymphoepithelial Kazal type-related inhibitor tends to be decreased in atopic dermatitis. J. Eur. Acad. Dermatol. Venereol. 23, 1263-1266
75. Roelandt, T., Thys, B., Heughebaert, C., De Vroede, A., De Paepe, K., Roseeuw, D., Rombaut, B., and Hachem, J.P. (2009). LEKTI-1 in sickness and in health. Int. J. Cosmet. Sci. 31, 247-254
76. Sato, J., Denda, M., Nakanishi, J., Nomura, J., and Koyama, J. (1998). Cholesterol sulfate inhibits proteases that are involved in desquamation of stratum corneum. J. Invest. Dermatol. 111, 189-193
77. Schalkwijk, J., de Roo, C., and de Jongh, G.J. (1991). Skin-derived antileukoproteinase (SKALP), an elastase inhibitor from human keratinocytes. Purification and biochemical properties. Biochim. Biophys. Acta 1096, 148-154
78. Schechter, N.M., Choi, E.J., Wang, Z.M., Hanakawa, Y., Stanley, J.R., Kang, Y., Clayman, G.L., and Jayakumar, A. (2005). Inhibition of human kallikreins 5 and 7 by the serine protease inhibitor lympho-epithelial Kazal-type inhibitor (LEKTI). Biol. Chem. 386, 1173-1184
79. Serre, G., Mils, V., Haftek, M., Vincent, C., Croute, F., Reano, A., Ouhayoun, J.P., Bettinger, S., and Soleilhavoup, J.P. (1991). Identification of late differentiation antigens of human cornified epithelia, expressed in re-organized desmosomes and bound to cross-linked envelope. J. Invest. Dermatol. 97, 1061-1072
80. Simon, M., Jonca, N., Guerrin, M., Haftek, M., Bernard, D., Caubet, C., Egelrud, T., Schmidt, R., and Serre, G. (2001). Refined characterization of corneodesmosin proteolysis during terminal differentiation of human epidermis and its relationship to desquamation. J. Biol. Chem. 276, 20292-20299
81. Sondell, B., Thornell, L.E., Stigbrand, T., and Egelrud, T. (1994). Immunolocalization of stratum corneum chymotryptic enzyme in human skin and oral epithelium with monoclonal antibodies: evidence of a proteinase specifically expressed in keratinizing squamous epithelia. J. Histochem. Cytochem. 42, 459-465
82. Sondell, B., Thornell, L.E., and Egelrud, T. (1995). Evidence that stratum corneum chymotryptic enzyme is transported to the stratum corneum extracellular space via lamellar bodies. J. Invest. Dermatol. 104, 819-823
83. Sondell, B., Dyberg, P., Anneroth, G.K., Ostman, P.O., and Egelrud, T. (1996). Association between expression of stratum corneum chymotryptic enzyme and pathological keratinization in human oral mucosa. Acta Derm. Venereol. 76, 177-181
84. Sprecher, E., Chavanas, S., DiGiovanna, J.J., Amin, S., Nielsen, K., Prendiville, J.S., Silverman, R., Esterly, N.B., Spraker, M.K., Guelig, E., de Luna, M.L., Williams, M.L., Buehler, B., Siegfried, E.C., van Maldergem, L., Pfendner, E., Bale, S.J., Uitto, J., Hovnanian, A., and Richard, G. (2001). The spectrum of pathogenic mutations in SPINK5 in 19 families with Netherton syndrome: implications for mutation detection and first case of prenatal diagnosis. J. Invest. Dermatol. 117, 179-187
85. Stefansson, K., Brattsand, M., Ny, A., Glas, B., and Egelrud, T. (2006). Kallikrein-related peptidase 14 may be a major contributor to trypsin-like proteolytic activity in human stratum corneum. Biol. Chem. 387, 761-768
86. Stefansson, K., Brattsand, M., Roosterman, D., Kempkes, C., Bocheva, G., Steinhoff, M., and Egelrud, T. (2008). Activation of proteinase-activated receptor-2 by human kallikrein-related peptidases. J. Invest. Dermatol. 128, 18-25
87. Suzuki, Y., Nomura, J., Hori, J., Koyama, J., Takahashi, M., and Horii, I. (1993). Detection and characterization of endogenous protease associated with desquamation of stratum corneum. Arch Dermatol. Res. 285, 372-377
88. Suzuki, Y., Nomura, J., Koyama, J., and Horii, I. (1994). The role of proteases in stratum corneum: involvement in stratum corneum desquamation. Arch Dermatol. Res. 286, 249-253
89. Suzuki, Y., Koyama, J., Moro, O., Horii, I., Kikuchi, K., Tanida, M., and Tagami, H. (1996). The role of two endogenous proteases of the stratum corneum in degradation of desmoglein-1 and their reduced activity in the skin of ichthyotic patients. Br. J. Dermatol. 134, 460-464
90. Tartaglia-Polcini, A., Bonnart, C., Micheloni, A., Cianfarani, F., Andre, A., Zambruno, G., Hovnanian, A., and D'Alessio, M. (2006). SPINK5, the defective gene in netherton syndrome, encodes multiple LEKTI isoforms derived from alternative pre-mRNA processing. J. Invest. Dermatol. 126, 315-324
91. Vasilopoulos, Y., Cork, M.J., Murphy, R., Williams, H.C., Robinson, D.A., Duff, G.W., Ward, S.J., and Tazi-Ahnini, R. (2004). Genetic association between an AACC insertion in the 3'UTR of the stratum corneum chymotryptic enzyme gene and atopic dermatitis. J. Invest. Dermatol. 123, 62-66
92. Wachter, A.M., and Lezdey, J. (1992). Treatment of atopic dermatitis with alpha 1-proteinase inhibitor. Ann. Allergy 69, 407-414
93. Walley, A.J., Chavanas, S., Moffatt, M.F., Esnouf, R.M., Ubhi, B., Lawrence, R., Wong, K., Abecasis, G.R., Jones, E.Y., Harper, J.I., Hovnanian, A., and Cookson, W.O. (2001). Gene polymorphism in Netherton and common atopic disease. Nat. Genet. 29, 175-178
94. Watkinson, A., Harding, C., Moore, A., and Coan, P. (2001). Water modulation of stratum corneum chymotryptic enzyme activity and desquamation. Arch Dermatol. Res. 293, 470-476
95. Wiedow, O., Schröder, J.M., Gregory, H., Young, J.A., and Christophers, E. (1990). Elafin: an elastasespecific inhibitor of human skin. Purification, characterization, and complete amino acid sequence. J. Biol. Chem. 265, 14791-14795
96. Wiedow, O., Harder, J., Bartels, J., Streit, V., and Christophers, E. (1998). Antileukoprotease in human skin: an antibiotic peptide constitutively produced by keratinocytes. Biochem. Biophys. Res. Commun. 248, 904-909
97. Wingens, M., van Bergen, B.H., Hiemstra, P.S., Meis, J.F., van Vlijmen-Willems, I.M., Zeeuwen, P.L., Mulder, J., Kramps, H.A., van Ruissen, F., and Schalkwijk, J. (1998). Induction of SLPI (ALP/ HUSI-I) in epidermal keratinocytes. J. Invest. Dermatol. 111, 996-1002
98. Wu, Z., Bartels, J., and Schroder, J.M. (2008). Lympho-Epithelial-Kazal-Type-Inhibitor (LEKTI)-2: a novel epithelial peptide antibiotic. Exp. Dermatol. 17, 260-260
99. Yamasaki, K., Schauber, J., Coda, A., Lin, H., Dorschner, R., Schechter, N., Bonnart, C., Descargues, P., Hovnanian, A., and Gallo, R. (2006). Kallikrein-mediated proteolysis regulates the antimicrobial effects of cathelicidins in skin. FASEB J. 20, 2068-2080
100. Yamasaki, K., Di Nardo, A., Bardan, A., Murakami, M., Ohtake, T., Coda, A., Dorschner, R.A., Bonnart, C., Descargues, P., Hovnanian, A., Morhenn, V.B., and Gallo, R.L. (2007). Increased serine protease activity and cathelicidin promotes skin inflammation in rosacea. Nat. Med. 13, 975-980
101. Yang, T., Liang, D., Koch, P.J., Hohl, D., Kheradmand, F., and Overbeek, P.A. (2004). Epidermal detachment, desmosomal dissociation, and destabilization of corneodesmosin in Spink5-/-mice. Genes Dev. 18, 2354-2358
102. Yoon, H., Laxmikanthan, G., Lee, J., Blaber, S.I., Rodriguez, A., Kogot, J.M., Scarisbrick, I.A., and Blaber, M. (2007). Activation profiles and regulatory cascades of the human kallikrein-related peptidases. J. Biol. Chem. 282, 31852-31864
103. Young, B., Heath, J.W., and Wheater, P.R., eds. (2000). Wheather's functional histology: a text and colour atlas, 4th edn (Churchill Livingstone: Edinburgh)