Stratum Corneum Defensive Functions: An Integrated View

Journal of Investigative Dermatology - Tập 125 Số 2 - Trang 183-200 - 2005
Peter M. Elias1,2
1Department of Dermatology, University of California, San Francisco, California, USA.
2Dermatology Service, Veterans Affairs Medical Center

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Alkemade, 1994, SKALP/elafin is an inducible proteinase inhibitor in human epidermal keratinocytes, J Cell Sci, 107, 2335, 10.1242/jcs.107.8.2335

Altemus, 2001, Stress-induced changes in skin barrier function in healthy women, J Invest Dermatol, 117, 309, 10.1046/j.1523-1747.2001.01373.x

Aly, 1978, Effect of prolonged occlusion on the microbial flora, pH, carbon dioxide and transepidermal water loss on human skin, J Invest Dermatol, 71, 378, 10.1111/1523-1747.ep12556778

Beare, 1958, The pH of the skin surface of children with seborrhoeic dermatitis compared with unaffected children, Br J Dermatol, 70, 233, 10.1111/j.1365-2133.1958.tb13330.x

Behne, 2001, The antiporter NHE1 influences the function of the SC pH gradient, J Skin Barrier Res, 3, 3

Behne, 2000, Omega-hydroxyceramides are required for corneocyte lipid envelope (CLE) formation and normal epidermal permeability barrier function, J Invest Dermatol, 114, 185, 10.1046/j.1523-1747.2000.00846.x

Behne, 2003, Neonatal development of the stratum corneum pH gradient: Localization and mechanisms leading to emergence of optimal barrier function, J Invest Dermatol, 120, 998, 10.1038/jid.2003.11

Behne, 2002, NHE1 regulates the stratum corneum permeability barrier homeostasis. Microenvironment acidification assessed with fluorescence lifetime imaging, J Biol Chem, 277, 47399, 10.1074/jbc.M204759200

Bernard, 2003, Analysis of proteins with caseinolytic activity in a human stratum corneum extract revealed a yet unidentified cysteine protease and identified the so-called “stratum corneum thiol protease” as cathepsin L2, J Invest Dermatol, 120, 592, 10.1046/j.1523-1747.2003.12086.x

Bibel, 1992, Antimicrobial activity of sphingosines, J Invest Dermatol, 98, 269, 10.1111/1523-1747.ep12497842

Bickenbach, 1995, Loricrin expression is coordinated with other epidermal proteins and the appearance of lipid lamellar granules in development, J Invest Dermatol, 104, 405, 10.1111/1523-1747.ep12665896

Bitoun, 2002, Netherton syndrome: Disease expression and spectrum of SPINK5 mutations in 21 families, J Invest Dermatol, 118, 352, 10.1046/j.1523-1747.2002.01603.x

Bouwstra, 2003, Water distribution and related morphology in human stratum corneum at different hydration levels, J Invest Dermatol, 120, 750, 10.1046/j.1523-1747.2003.12128.x

Bouwstra, 1998, pH, cholesterol sulfate, and fatty acids affect the stratum corneum lipid organization, J Invest Dermatol Symp Proc, 3, 69, 10.1038/jidsymp.1998.17

Bouwstra, 2000, Phase behaviour of skin barrier model membranes at pH 7.4, Cell Mol Biol, 46, 979

Braff, 2005, Keratinocytes store the antimicrobial peptide cathelicidin in lamellar bodies, J Invest Dermatol, 124, 394, 10.1111/j.0022-202X.2004.23443.x

Braissant, 1996, Differential expression of peroxisome proliferator-activated receptors (PPARs): Tissue distribution of PPAR-alpha, -beta, and -gamma in the adult rat, Endocrinology, 137, 354, 10.1210/endo.137.1.8536636

Braissant, 1998, Differential expression of peroxisome proliferator-activated receptor-alpha, -beta, and -gamma during rat embryonic development, Endocrinology, 139, 2748, 10.1210/endo.139.6.6049

Brattsand, 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, 10.1074/jbc.274.42.30033

Brown, 1998, Sterol regulatory element binding proteins (SREBPs): Controllers of lipid synthesis and cellular uptake, Nutr Rev, 56, 1, 10.1111/j.1753-4887.1998.tb01680.x

Candi, 1998, A highly conserved lysine residue on the head domain of type II keratins is essential for the attachment of keratin intermediate filaments to the cornified cell envelope through isopeptide crosslinking by transglutaminases, Proc Natl Acad Sci USA, 95, 2067, 10.1073/pnas.95.5.2067

Caubet, 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, 10.1111/j.0022-202X.2004.22512.x

Chakravarty, 1989, Acylation of keratinocyte transglutaminase by palmitic and myristic acids in the membrane anchorage region, J Biol Chem, 264, 625, 10.1016/S0021-9258(17)31306-6

Chamlin, 2002, Ceramide-dominant barrier repair lipids alleviate childhood atopic dermatitis: Changes in barrier function provide a sensitive indicator of disease activity, J Am Acad Dermatol, 47, 198, 10.1067/mjd.2002.124617

Chapman, 1991, Lipids, proteins and corneocyte adhesion, Arch Dermatol Res, 283, 167, 10.1007/BF00372057

Chavanas, 2000, Mutations in SPINK5, encoding a serine protease inhibitor, cause Netherton syndrome, Nat Genet, 25, 141, 10.1038/75977

Choate, 1996, Transglutaminase 1 delivery to lamellar ichthyosis keratinocytes, Hum Gene Ther, 7, 2247, 10.1089/hum.1996.7.18-2247

Choate, 1996, Corrective gene transfer in the human skin disorder lamellar ichthyosis, Nat Med, 2, 1263, 10.1038/nm1196-1263

Choate, 1998, Transglutaminase 1 expression in a patient with features of harlequin ichthyosis: Case report, J Am Acad Dermatol, 38, 325, 10.1016/S0190-9622(98)70575-4

Christophers, 1964, Visualization of the cell layers of the stratum corneum, J Invest Dermatol, 42, 407, 10.1038/jid.1964.88

Chujor, 1998, Glucosylceramide synthase activity in murine epidermis: Quantitation, localization, regulation, and requirement for barrier homeostasis, J Lipid Res, 39, 277, 10.1016/S0022-2275(20)33889-X

Corden, 1996, Human keratin diseases: Hereditary fragility of specific epithelial tissues, Exp Dermatol, 5, 297, 10.1111/j.1600-0625.1996.tb00133.x

Dale, 1997, Transient expression of epidermal filaggrin in cultured cells causes collapse of intermediate filament networks with alteration of cell shape and nuclear integrity, J Invest Dermatol, 108, 179, 10.1111/1523-1747.ep12334205

Denda, 1998, Exposure to a dry environment enhances epidermal permeability barrier function, J Invest Dermatol, 111, 858, 10.1046/j.1523-1747.1998.00333.x

Denda, 2000, Stress alters cutaneous permeability barrier homeostasis, Am J Physiol Regul Integr Comp Physiol, 278, 367, 10.1152/ajpregu.2000.278.2.R367

Denda, 1996, The epidermal hyperplasia associated with repeated barrier disruption by acetone treatment or tape stripping cannot be attributed to increased water loss, Arch Dermatol Res, 288, 230, 10.1007/BF02530090

De Simone, 2001, Urocanic acid isomers in patients with non-melanoma skin cancer, Br J Dermatol, 144, 858, 10.1046/j.1365-2133.2001.04145.x

Doering, 1999, Sphingolipid activator proteins are required for epidermal permeability barrier formation, J Biol Chem, 274, 11038, 10.1074/jbc.274.16.11038

Dunsche, 2002, The novel human beta-defensin-3 is widely expressed in oral tissues, Eur J Oral Sci, 110, 121, 10.1034/j.1600-0722.2002.11186.x

Duplus, 2002, Is there a single mechanism for fatty acid regulation of gene transcription?, Biochem Pharmacol, 64, 893, 10.1016/S0006-2952(02)01157-7

Eckert, 1997, The epidermal keratinocyte as a model for the study of gene regulation and cell differentiation, Physiol Rev, 77, 397, 10.1152/physrev.1997.77.2.397

Egberts, 2004, Cathepsin D is involved in the regulation of transglutaminase 1 and epidermal differentiation, J Cell Sci, 117, 2295, 10.1242/jcs.01075

Ekanayake-Mudiyanselage, 1998, Proksch E: Expression of epidermal keratins and the cornified envelope protein involucrin is influenced by permeability barrier disruption, J Invest Dermatol, 111, 517, 10.1046/j.1523-1747.1998.00318.x

Ekholm, 2000, Stratum corneum tryptic enzyme in normal epidermis: A missing link in the desquamation process?, J Invest Dermatol, 114, 56, 10.1046/j.1523-1747.2000.00820.x

Elias, 2002, Origin of the epidermal calcium gradient: Regulation by barrier status and role of active vs passive mechanisms, J Invest Dermatol, 119, 1269, 10.1046/j.1523-1747.2002.19622.x

Elias, 2002, Modulations in epidermal calcium regulate the expression of differentiation-specific markers, J Invest Dermatol, 119, 1128, 10.1046/j.1523-1747.2002.19512.x

Elias, 1978, The mammalian cutaneous permeability barrier: Defective barrier function is essential fatty acid deficiency correlates with abnormal intercellular lipid deposition, Lab Invest, 39, 574

Elias, 1980, The permeability barrier in essential fatty acid deficiency: Evidence for a direct role for linoleic acid in barrier function, J Invest Dermatol, 74, 230, 10.1111/1523-1747.ep12541775

Elias, 2004, Basis for abnormal desquamation and permeability barrier dysfunction in RXLI, J Invest Dermatol, 122, 314, 10.1046/j.1523-1747.2003.22258.x

Elias, 1998, The secretory granular cell: The outermost granular cell as a specialized secretory cell, J Invest Dermatol Symp Proc, 3, 87, 10.1038/jidsymp.1998.20

Elias, 2000, Origin of the corneocyte lipid envelope (CLE): Observations in harlequin ichthyosis and cultured human keratinocytes, J Invest Dermatol, 115, 765, 10.1046/j.1523-1747.2000.00124-5.x

Elias, 1992, Lipids and the epidermal water barrier: Metabolism, regulation, and pathophysiology, Semin Dermatol, 11, 176

Elias, 1999, Skin as an organ of protection, 164

Elias, 2001, Does the tail wag the dog? Role of the barrier in the pathogenesis of inflammatory dermatoses and therapeutic implications, Arch Dermatol, 137, 1079

Elias, 2003, Skin as an organ of protection, 164

Elias, 2002, The aged epidermal permeability barrier: Basis for functional abnormalities, Clin Geriatr Med, 18, 103, 10.1016/S0749-0690(03)00037-5

Elias, 1977, Mammalian epidermal barrier layer lipids: Composition and influence on structure, J Invest Dermatol, 69, 535, 10.1111/1523-1747.ep12687968

Elias, 1977, Membrane alterations during cornification of mammalian squamous epithelia: A freeze-fracture, tracer, and thin-section study, Anat Rec, 189, 577, 10.1002/ar.1091890404

Elias, 1991, Structural and lipid biochemical correlates of the epidermal permeability barrier, Adv Lipid Res, 24, 1, 10.1016/B978-0-12-024924-4.50005-5

Elias, 1988, Membrane structural alterations in murine stratum corneum: Relationship to the localization of polar lipids and phospholipases, J Invest Dermatol, 91, 3, 10.1111/1523-1747.ep12463279

Elias, 2002, Basis for the permeability barrier abnormality in lamellar ichthyosis, Exp Dermatol, 11, 248, 10.1034/j.1600-0625.2001.110308.x

Elias, 1999, Epidermal pathogenesis of inflammatory dermatoses, Am J Contact Dermat, 10, 119

Ellis, 2000, Troglitazone improves psoriasis and normalizes models of proliferative skin disease: Ligands for peroxisome proliferator-activated receptor-gamma inhibit keratinocyte proliferation, Arch Dermatol, 136, 609, 10.1001/archderm.136.5.609

Farrell, 1998, UVB irradiation up-regulates serine palmitoyltransferase in cultured human keratinocytes, J Lipid Res, 39, 2031, 10.1016/S0022-2275(20)32501-3

Fartasch, 1999, Altered lamellar body secretion and stratum corneum membrane structure in Netherton syndrome: Differentiation from other infantile erythrodermas and pathogenic implications, Arch Dermatol, 135, 823, 10.1001/archderm.135.7.823

Feingold, 1991, The regulation and role of epidermal lipid synthesis, Adv Lipid Res, 24, 57, 10.1016/B978-0-12-024924-4.50007-9

Feingold, 1983, Localization of de novo sterologenesis in mammalian skin, J Invest Dermatol, 81, 365, 10.1111/1523-1747.ep12519974

Feingold, 1990, Cholesterol synthesis is required for cutaneous barrier function in mice, J Clin Invest, 86, 1738, 10.1172/JCI114899

Feingold, 1991, The lovastatin-treated rodent: A new model of barrier disruption and epidermal hyperplasia, J Invest Dermatol, 96, 201, 10.1111/1523-1747.ep12461153

Feliciani, 1996, Keratinocytes and cytokine/growth factors, Crit Rev Oral Biol Med, 7, 300, 10.1177/10454411960070040101

Finlay-Jones, 1998, Photoprotection: Sunscreens and the immunomodulatory effects of UV irradiation, Mutat Res, 422, 155, 10.1016/S0027-5107(98)00187-0

Fitzgerald, 2002, Nuclear hormone receptors and cholesterol trafficking: The orphans find a new home, J Mol Med, 80, 271, 10.1007/s00109-001-0318-y

Fleckman, 1985, Profilaggrin, a high-molecular-weight precursor of filaggrin in human epidermis and cultured keratinocytes, J Invest Dermatol, 85, 507, 10.1111/1523-1747.ep12277306

Fluhr, 2004, Stratum corneum acidification in neonatal skin: Secretory phospholipase A2 and the sodium/hydrogen antiporter-1 acidify neonatal rat stratum corneum, J Invest Dermatol, 122, 320, 10.1046/j.0022-202X.2003.00204.x

Fluhr, 2001, Generation of free fatty acids from phospholipids regulates stratum corneum acidification and integrity, J Invest Dermatol, 117, 44, 10.1046/j.0022-202x.2001.01399.x

Fluhr, 2003, Glycerol regulates stratum corneum hydration in sebaceous gland deficient (asebia) mice, J Invest Dermatol, 120, 728, 10.1046/j.1523-1747.2003.12134.x

Fowler, 2003, Liver X receptor activators display anti-inflammatory activity in irritant and allergic contact dermatitis models: Liver-X-receptor-specific inhibition of inflammation and primary cytokine production, J Invest Dermatol, 120, 246, 10.1046/j.1523-1747.2003.12033.x

Freinkel, 1983, Acid hydrolases of the epidermis: Subcellular localization and relationship to cornification, J Invest Dermatol, 80, 441, 10.1111/1523-1747.ep12555534

Fuchs, 1994, Intermediate filaments and disease: Mutations that cripple cell strength, J Cell Biol, 125, 511, 10.1083/jcb.125.3.511

Fuchs, 1994, Intermediate filaments: Structure, dynamics, function, and disease, Annu Rev Biochem, 63, 345, 10.1146/annurev.bi.63.070194.002021

Gallo, 2003, Endogenous production of antimicrobial peptides in innate immunity and human disease, Curr Allergy Asthma Rep, 3, 402, 10.1007/s11882-003-0074-x

Ganz, 2003, Defensins: Antimicrobial peptides of innate immunity, Nat Rev Immunol, 3, 710, 10.1038/nri1180

Garg, 2001, Psychological stress perturbs epidermal permeability barrier homeostasis: Implications for the pathogenesis of stress-associated skin disorders, Arch Dermatol, 137, 53, 10.1001/archderm.137.1.53

Ghadially, 1996, Decreased epidermal lipid synthesis accounts for altered barrier function in aged mice, J Invest Dermatol, 106, 1064, 10.1111/1523-1747.ep12338692

Ghadially, 1995, The aged epidermal permeability barrier. Structural, functional, and lipid biochemical abnormalities in humans and a senescent murine model, J Clin Invest, 95, 2281, 10.1172/JCI117919

Ghadially, 1992, Effects of petrolatum on stratum corneum structure and function, J Am Acad Dermatol, 26, 387, 10.1016/0190-9622(92)70060-S

Ghadially, 1996, Stratum corneum structure and function correlates with phenotype in psoriasis, J Invest Dermatol, 107, 558, 10.1111/1523-1747.ep12582813

Ghadially, 1992, Membrane structural abnormalities in the stratum corneum of the autosomal recessive ichthyoses, J Invest Dermatol, 99, 755, 10.1111/1523-1747.ep12614489

Giusti, 2001, Skin barrier, hydration, and pH of the skin of infants under 2 years of age, Pediatr Dermatol, 18, 93, 10.1046/j.1525-1470.2001.018002093.x

Gläser, 2005, Antimicrobial psoriasin (S100A7) protects human skin from Escherichia coli infection, Nat Immunol, 6, 57, 10.1038/ni1142

Grayson, 1985, Lamellar body-enriched fractions from neonatal mice: Preparative techniques and partial characterization, J Invest Dermatol, 85, 289, 10.1111/1523-1747.ep12276826

Grubauer, 1989, Transepidermal water loss: The signal for recovery of barrier structure and function, J Lipid Res, 30, 323, 10.1016/S0022-2275(20)38361-9

Grubauer, 1987, Relationship of epidermal lipogenesis to cutaneous barrier function, J Lipid Res, 28, 746, 10.1016/S0022-2275(20)38670-3

Hachem, 2003, pH directly regulates epidermal permeability barrier homeostasis, and stratum corneum integrity/cohesion, J Invest Dermatol, 121, 345, 10.1046/j.1523-1747.2003.12365.x

Haftek, 1996, A longitudinal study of a harlequin infant presenting clinically as non-bullous congenital ichthyosiform erythroderma, Br J Dermatol, 135, 448, 10.1111/j.1365-2133.1996.tb01512.x

Hanley, 1998, Glucocorticoid deficiency delays stratum corneum maturation in the fetal mouse, J Invest Dermatol, 111, 440, 10.1046/j.1523-1747.1998.00303.x

Hanley, 1997, Activators of the nuclear hormone receptors PPARalpha and FXR accelerate the development of the fetal epidermal permeability barrier, J Clin Invest, 100, 705, 10.1172/JCI119583

Hanley, 1999, Fetal epidermal differentiation and barrier development in vivo is accelerated by nuclear hormone receptor activators, J Invest Dermatol, 113, 788, 10.1046/j.1523-1747.1999.00743.x

Hanley, 2000, Farnesol stimulates differentiation in epidermal keratinocytes via PPARalpha, J Biol Chem, 275, 11484, 10.1074/jbc.275.15.11484

Hanley, 2000, Oxysterols induce differentiation in human keratinocytes and increase Ap-1-dependent involucrin transcription, J Invest Dermatol, 114, 545, 10.1046/j.1523-1747.2000.00895.x

Hanley, 1996, Epidermal barrier ontogenesis: Maturation in serum-free media and acceleration by glucocorticoids and thyroid hormone but not selected growth factors, J Invest Dermatol, 106, 404, 10.1111/1523-1747.ep12343405

Hanley, 1996, Hormonal basis for the gender difference in epidermal barrier formation in the fetal rat. Acceleration by estrogen and delay by testosterone, J Clin Invest, 97, 2576, 10.1172/JCI118706

Hanson, 2002, Two-photon fluorescence lifetime imaging of the skin stratum corneum pH gradient, Biophys J, 83, 1682, 10.1016/S0006-3495(02)73936-2

Hansson, 1994, Cloning, expression, and characterization of stratum corneum chymotryptic enzyme. A skin-specific human serine proteinase, J Biol Chem, 269, 19420, 10.1016/S0021-9258(17)32185-3

Hara, 2000, High-expression of sphingomyelin deacylase is an important determinant of ceramide deficiency leading to barrier disruption in atopic dermatitis, J Invest Dermatol, 115, 406, 10.1046/j.1523-1747.2000.00072.x

Harder, 2002, RNase 7, a novel innate immune defense antimicrobial protein of healthy human skin, J Biol Chem, 277, 46779, 10.1074/jbc.M207587200

Harder, 2005, Psoriatic scales: A promising source for the isolation of human skin-derived antimicrobial proteins, J Leukoc Biol, 77, 476, 10.1189/jlb.0704409

Harding, 2000, Dry skin, moisturization, and corneodesmolysis, Inter J Cosm Sci, 22, 21, 10.1046/j.1467-2494.2000.00001.x

Harris, 1997, Permeability barrier disruption coordinately regulates mRNA levels for key enzymes of cholesterol, fatty acid, and ceramide synthesis in the epidermis, J Invest Dermatol, 109, 783, 10.1111/1523-1747.ep12340962

Harris, 1998, Parallel regulation of sterol regulatory element binding protein-2 and the enzymes of cholesterol and fatty acid synthesis but not ceramide synthesis in cultured human keratinocytes and murine epidermis, J Lipid Res, 39, 412, 10.1016/S0022-2275(20)33902-X

Hennings, 1981, Calcium induction of transglutaminase and the formation of epsilon (gamma-glutamyl) lysine cross-links in cultured mouse epidermal cells, Biochem Biophys Res Commun, 102, 739, 10.1016/S0006-291X(81)80194-5

Hohl, 1990, Cornified cell envelope, Dermatologica, 180, 201, 10.1159/000248031

Hohl, 1993, Analysis of the cornified cell envelope in lamellar ichthyosis, Arch Dermatol, 129, 618, 10.1001/archderm.1993.01680260088013

Hohl, 1991, Characterization of human loricrin. Structure and function of a new class of epidermal cell envelope proteins, J Biol Chem, 266, 6626, 10.1016/S0021-9258(18)38163-8

Holleran, 1991, Regulation of epidermal sphingolipid synthesis by permeability barrier function, J Lipid Res, 32, 1151, 10.1016/S0022-2275(20)41978-9

Holleran, 1991, Sphingolipids are required for mammalian epidermal barrier function. Inhibition of sphingolipid synthesis delays barrier recovery after acute perturbation, J Clin Invest, 88, 1338, 10.1172/JCI115439

Horikoshi, 1998, Isoforms of cathepsin D and human epidermal differentiation, Biochimie, 80, 605, 10.1016/S0300-9084(98)80013-8

Horikoshi, 1999, Role of endogenous cathepsin D-like and chymotrypsin-like proteolysis in human epidermal desquamation, Br J Dermatol, 141, 453, 10.1046/j.1365-2133.1999.03038.x

Howell, 2004, Selective killing of vaccinia virus by LL-37: Implications for eczema vaccinatum, J Immunol, 172, 1763, 10.4049/jimmunol.172.3.1763

Huber, 1995, Mutations of keratinocyte transglutaminase in lamellar ichthyosis, Science, 267, 525, 10.1126/science.7824952

Huh, 2002, Dynamic alteration of human beta-defensin 2 localization from cytoplasm to intercellular space in psoriatic skin, J Mol Med, 80, 678, 10.1007/s00109-002-0373-z

Hurt, 1995, Cutaneous lipid synthesis during late fetal development in the rat, Arch Dermatol Res, 287, 754, 10.1007/BF01105801

Imokawa, 1991, Decreased level of ceramides in stratum corneum of atopic dermatitis: An etiologic factor in atopic dry skin?, J Invest Dermatol, 96, 523, 10.1111/1523-1747.ep12470233

Ishida-Yamamoto, 1998, Structural organization of cornified cell envelopes and alterations in inherited skin disorders, Exp Dermatol, 7, 1, 10.1111/j.1600-0625.1998.tb00295.x

Jensen, 1999, Roles for tumor necrosis factor receptor p55 and sphingomyelinase in repairing the cutaneous permeability barrier, J Clin Invest, 104, 1761, 10.1172/JCI5307

Jonca, 2002, Corneodesmosin, a component of epidermal corneocyte desmosomes, displays homophilic adhesive properties, J Biol Chem, 277, 5024, 10.1074/jbc.M108438200

Kalinin, 2002, Epithelial barrier function: Assembly and structural features of the cornified cell envelope, Bioessays, 24, 789, 10.1002/bies.10144

Kao, 2003, Short-term glucocorticoid treatment compromises both permeability barrier homeostasis and stratum corneum integrity: Inhibition of epidermal lipid synthesis accounts for functional abnormalities, J Invest Dermatol, 120, 456, 10.1046/j.1523-1747.2003.12053.x

Kao, 2001, Testosterone perturbs epidermal permeability barrier homeostasis, J Invest Dermatol, 116, 443, 10.1046/j.1523-1747.2001.01281.x

Kimonis, 1994, A mutation in the V1 end domain of keratin 1 in non-epidermolytic palmar-plantar keratoderma, J Invest Dermatol, 103, 764, 10.1111/1523-1747.ep12412771

Komatsu, 2002, Elevated stratum corneum hydrolytic activity in Netherton syndrome suggests an inhibitory regulation of desquamation by SPINK5-derived peptides, J Invest Dermatol, 118, 436, 10.1046/j.0022-202x.2001.01663.x

Komuves, 1998, Ligands and activators of nuclear hormone receptors regulate epidermal differentiation during fetal rat skin development, J Invest Dermatol, 111, 429, 10.1046/j.1523-1747.1998.00296.x

Komuves, 2000, Keratinocyte differentiation in hyperproliferative epidermis: Topical application of PPARalpha activators restores tissue homeostasis, J Invest Dermatol, 115, 361, 10.1046/j.1523-1747.2000.00076.x

Korting, 1990, Differences in the skin surface pH and bacterial microflora due to the long-term application of synthetic detergent preparations of pH 5.5 and pH 7.0. Results of a crossover trial in healthy volunteers, Acta Derm Venereol, 70, 429, 10.2340/0001555570429431

Kreder, 1999, Impaired neutral sphingomyelinase activation and cutaneous barrier repair in FAN-deficient mice, EMBO J, 18, 2472, 10.1093/emboj/18.9.2472

Krien, 2000, Evidence for the existence of a self-regulated enzymatic process within human stratum corneum—an unexpected role for urocanic acid, J Invest Dermatol, 115, 414, 10.1046/j.1523-1747.2000.00083.x

Kupper, 1990, Immune and inflammatory processes in cutaneous tissues. Mechanisms and speculations, J Clin Invest, 86, 1783, 10.1172/JCI114907

Lampe, 1983, Human stratum corneum lipids: Characterization and regional variations, J Lipid Res, 24, 120, 10.1016/S0022-2275(20)38005-6

Landmann, 1988, The epidermal permeability barrier, Anat Embryol (Berl), 178, 1, 10.1007/BF00305008

Lavrijsen, 1995, Reduced skin barrier function parallels abnormal stratum corneum lipid organization in patients with lamellar ichthyosis, J Invest Dermatol, 105, 619, 10.1111/1523-1747.ep12323752

Lee, 1992, Calcium and potassium are important regulators of barrier homeostasis in murine epidermis, J Clin Invest, 89, 530, 10.1172/JCI115617

Leyden, 1979, Updated in vivo methods for evaluating topical antimicrobial agents on human skin, J Invest Dermatol, 72, 165, 10.1111/1523-1747.ep12676347

Liou, 1997, Amphiregulin and nerve growth factor expression are regulated by barrier status in murine epidermis, J Invest Dermatol, 108, 73, 10.1111/1523-1747.ep12285638

Liu, 2002, Human beta-defensin-2 production in keratinocytes is regulated by interleukin-1, bacteria, and the state of differentiation, J Invest Dermatol, 118, 275, 10.1046/j.0022-202x.2001.01651.x

Loewen, 2002, Cholesterol homeostasis: Not until the SCAP lady INSIGs, Curr Biol, 12, R779, 10.1016/S0960-9822(02)01292-7

Lundstrom, 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, 10.1007/BF00371795

Madison, 1998, Lamellar granule biogenesis: A role for ceramide glucosyltransferase, lysosomal enzyme transport, and the Golgi, J Invest Dermatol Symp Proc, 3, 80, 10.1038/jidsymp.1998.19

Maestrini, 1996, A molecular defect in loricrin, the major component of the cornified cell envelope, underlies Vohwinkel's syndrome, Nat Genet, 13, 70, 10.1038/ng0596-70

Man, 1996, Optimization of physiological lipid mixtures for barrier repair, J Invest Dermatol, 106, 1096, 10.1111/1523-1747.ep12340135

Man, 1993, Exogenous lipids influence permeability barrier recovery in acetone-treated murine skin, Arch Dermatol, 129, 728, 10.1001/archderm.1993.01680270066008

Mao-Qiang, 1995, Exogenous nonphysiologic vs physiologic lipids. Divergent mechanisms for correction of permeability barrier dysfunction, Arch Dermatol, 131, 809, 10.1001/archderm.1995.01690190063012

Mao-Qiang, 1993, Fatty acids are required for epidermal permeability barrier function, J Clin Invest, 92, 791, 10.1172/JCI116652

Mao-Qiang, 1995, Extracellular processing of phospholipids is required for permeability barrier homeostasis, J Lipid Res, 36, 1925, 10.1016/S0022-2275(20)41111-3

Mao-Qiang, 1996, Secretory phospholipase A2 activity is required for permeability barrier homeostasis, J Invest Dermatol, 106, 57, 10.1111/1523-1747.ep12327246

Marekov, 1998, Ceramides are bound to structural proteins of the human foreskin epidermal cornified cell envelope, J Biol Chem, 273, 17763, 10.1074/jbc.273.28.17763

Mauro, 1998, Barrier recovery is impeded at neutral pH, independent of ionic effects: Implications for extracellular lipid processing, Arch Dermatol Res, 290, 215, 10.1007/s004030050293

Maury, 2000, Identification of two secreted phospholipases A2 in human epidermis, J Invest Dermatol, 114, 960, 10.1046/j.1523-1747.2000.00965.x

Mazereeuw-Hautier, 2000, Identification of pancreatic type I secreted phospholipase A2 in human epidermis and its determination by tape stripping, Br J Dermatol, 142, 424, 10.1046/j.1365-2133.2000.03351.x

Menon, 1997, Morphologic basis for a pore-pathway in mammalian stratum corneum, Skin Pharmacol, 10, 235, 10.1159/000211511

Menon, 1994, Integrity of the permeability barrier is crucial for maintenance of the epidermal calcium gradient, Br J Dermatol, 130, 139, 10.1111/j.1365-2133.1994.tb02892.x

Menon, 1992, Lamellar body secretory response to barrier disruption, J Invest Dermatol, 98, 279, 10.1111/1523-1747.ep12497866

Menon, 1985, De novo sterologenesis in the skin. II. Regulation by cutaneous barrier requirements, J Lipid Res, 26, 418, 10.1016/S0022-2275(20)34355-8

Menon, 1985, Ionic calcium reservoirs in mammalian epidermis: Ultrastructural localization by ion-capture cytochemistry, J Invest Dermatol, 84, 508, 10.1111/1523-1747.ep12273485

Menon, 1994, Selective obliteration of the epidermal calcium gradient leads to enhanced lamellar body secretion, J Invest Dermatol, 102, 789, 10.1111/1523-1747.ep12377921

Menton, 1971, Structure and organization of mammalian stratum corneum, J Ultrastruct Res, 35, 247, 10.1016/S0022-5320(71)80155-7

Miller, 1988, In vitro and in vivo antistaphylococcal activity of human stratum corneum lipids, Arch Dermatol, 124, 209, 10.1001/archderm.1988.01670020027012

Molhuizen, 1993, SKALP/elafin: An elastase inhibitor from cultured human keratinocytes. Purification, cDNA sequence, and evidence for transglutaminase cross-linking, J Biol Chem, 268, 12028, 10.1016/S0021-9258(19)50303-9

Moskowitz DG, Fowler AJ, Heyman MB, Cohen SP, Crumrine D, Elias PM, Williams ML: Pathophysiological basis for growth failure in children with ichthyosis: An evaluation of cutaneous ultrastructure, epidermal permeability barrier function, and energy expenditure. J Pediatr, in press

Narce, 2002, Lipid metabolism: Regulation of lipid metabolism gene expression by peroxisome proliferator-activated receptor alpha and sterol regulatory element binding proteins, Curr Opin Lipidol, 13, 445, 10.1097/00041433-200208000-00013

Nickoloff, 1994, Perturbation of epidermal barrier function correlates with initiation of cytokine cascade in human skin, J Am Acad Dermatol, 30, 535, 10.1016/S0190-9622(94)70059-1

Nizet, 2001, Innate antimicrobial peptide protects the skin from invasive bacterial infection, Nature, 414, 454, 10.1038/35106587

Noonan, 1992, Immunosuppression by ultraviolet B radiation: Initiation by urocanic acid, Immunol Today, 13, 250, 10.1016/0167-5699(92)90005-R

Norlen, 2001, Skin barrier formation: The membrane folding model, J Invest Dermatol, 117, 823, 10.1046/j.0022-202x.2001.01445.x

Norlen, 2001, Skin barrier structure and function: The single gel phase model, J Invest Dermatol, 117, 830, 10.1038/jid.2001.1

Odland, 1981, The lamellar granules of epidermis, Curr Probl Dermatol, 9, 29, 10.1159/000403343

Ohman, 1994, In vivo studies concerning a pH gradient in human stratum corneum and upper epidermis, Acta Derm Venereol, 74, 375, 10.2340/0001555574375379

Ong, 2002, Endogenous antimicrobial peptides and skin infections in atopic dermatitis, N Engl J Med, 347, 1151, 10.1056/NEJMoa021481

Oren, 2003, In human epidermis, beta-defensin 2 is packaged in lamellar bodies, Exp Mol Pathol, 74, 180, 10.1016/S0014-4800(02)00023-0

Orlowski, 1997, Na+/H+ exchangers of mammalian cells, J Biol Chem, 272, 22373, 10.1074/jbc.272.36.22373

Ottey, 1995, Cutaneous permeability barrier disruption increases fatty acid synthetic enzyme activity in the epidermis of hairless mice, J Invest Dermatol, 104, 401, 10.1111/1523-1747.ep12665893

Potts, 1990, Lipid biophysics of water loss through the skin, Proc Natl Acad Sci USA, 87, 3871, 10.1073/pnas.87.10.3871

Proksch, 1990, Regulation of 3-hydroxy-3-methylglutaryl-coenzyme A reductase activity in murine epidermis. Modulation of enzyme content and activation state by barrier requirements, J Clin Invest, 85, 874, 10.1172/JCI114514

Proksch, 2003, Skin lipids and epidermal differentiation in atopic dermatitis, Clin Dermatol, 21, 134, 10.1016/S0738-081X(02)00370-X

Rawson, 2003, The SREBP pathway—insights from Insigs and insects, Nat Rev Mol Cell Biol, 4, 631, 10.1038/nrm1174

Redoules, 1999, Characterisation and assay of five enzymatic activities in the stratum corneum using tape-strippings, Skin Pharmacol Appl Skin Physiol, 12, 182, 10.1159/000066242

Resing, 1993, Independent regulation of two cytoplasmic processing stages of the intermediate filament-associated protein filaggrin and role of Ca2+ in the second stage, J Biol Chem, 268, 25139, 10.1016/S0021-9258(19)74580-3

Risse, 1998, Differentiating cells of murine stratified squamous epithelia constitutively express plasminogen activator inhibitor type 2 (PAI-2), Histochem Cell Biol, 110, 559, 10.1007/s004180050318

Risse, 2000, Evidence for intracellular cleavage of plasminogen activator inhibitor type 2 (PAI-2) in normal epidermal keratinocytes, J Cell Physiol, 182, 281, 10.1002/(SICI)1097-4652(200002)182:2<281::AID-JCP17>3.0.CO;2-D

Rivier, 2000, Peroxisome proliferator-activated receptor-alpha enhances lipid metabolism in a skin equivalent model, J Invest Dermatol, 114, 681, 10.1046/j.1523-1747.2000.00939.x

Rothnagel, 1992, Mutations in the rod domains of keratins 1 and 10 in epidermolytic hyperkeratosis, Science, 257, 1128, 10.1126/science.257.5073.1128

Ruhl, 2003, Inhibition of IgE-production by peroxisome proliferator-activated receptor ligands, J Invest Dermatol, 121, 757, 10.1046/j.1523-1747.2003.12493.x

Sarangarajan, 2001, Molecular and functional characterization of sodium–hydrogen exchanger in skin as well as cultured keratinocytes and melanocytes, Biochim Biophys Acta, 1511, 181, 10.1016/S0005-2736(01)00273-5

Sato, 2002, Abrupt decreases in environmental humidity induce abnormalities in permeability barrier homeostasis, J Invest Dermatol, 119, 900, 10.1046/j.1523-1747.2002.00589.x

Schadow, 2001, Characterization of group X phospholipase A(2) as the major enzyme secreted by human keratinocytes and its regulation by the phorbol ester TPA, J Invest Dermatol, 116, 31, 10.1046/j.1523-1747.2001.00179.x

Scheuplein, 1971, Permeability of the skin, Physiol Rev, 51, 702, 10.1152/physrev.1971.51.4.702

Schmuth, 2004, The effect of LXR activators on AP-1 proteins in keratinocytes, J Invest Dermatol, 123, 41, 10.1111/j.0022-202X.2004.22707.x

Schmuth, 2004, Structural and functional consequences of loricrin mutations in human loricrin keratoderma (Vohwinkel syndrome with ichthyosis), J Invest Dermatol, 122, 909, 10.1111/j.0022-202X.2004.22431.x

Schmuth, 2004, Peroxisome proliferator-activated receptor (PPAR)-beta/delta stimulates differentiation and lipid accumulation in keratinocytes, J Invest Dermatol, 122, 971, 10.1111/j.0022-202X.2004.22412.x

Schmuth, 2001, Pathogenesis of the permeability barrier abnormality in epidermolytic hyperkeratosis, J Invest Dermatol, 117, 837, 10.1046/j.0022-202x.2001.01471.x

Schurer, 1991, The biochemistry and function of stratum corneum lipids, Adv Lipid Res, 24, 27, 10.1016/B978-0-12-024924-4.50006-7

Scott, 1981, Factors controlling the expressed activity of histidine ammonia-lyase in the epidermis and the resulting accumulation of urocanic acid, Biochem J, 194, 829, 10.1042/bj1940829

Scott, 1986, Filaggrin breakdown to water binding compounds during development of the rat stratum corneum is controlled by the water activity of the environment, Dev Biol, 115, 84, 10.1016/0012-1606(86)90230-7

Serre, 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, 10.1111/1523-1747.ep12492589

Sheu, 2002, Topical peroxisome proliferator activated receptor-alpha activators reduce inflammation in irritant and allergic contact dermatitis models, J Invest Dermatol, 118, 94, 10.1046/j.0022-202x.2001.01626.x

Simon, 2001, Refined characterization of corneodesmosin proteolysis during terminal differentiation of human epidermis and its relationship to desquamation, J Biol Chem, 276, 20292, 10.1074/jbc.M100201200

Simon, 1997, Characterization and purification of human corneodesmosin, an epidermal basic glycoprotein associated with corneocyte-specific modified desmosomes, J Biol Chem, 272, 31770, 10.1074/jbc.272.50.31770

Smith, 1988, Multiple sterol regulatory elements in promoter for hamster 3-hydroxy-3-methylglutaryl-coenzyme A synthase, J Biol Chem, 263, 18480, 10.1016/S0021-9258(19)81383-2

Sondell, 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, 10.1177/42.4.7510318

Sparavigna, 1999, Cutaneous pH in children affected by atopic dermatitis and in healthy children: A multicenter study, Skin Res Technol, 5, 221, 10.1111/j.1600-0846.1999.tb00134.x

Steinert, 1997, Direct evidence that involucrin is a major early isopeptide cross-linked component of the keratinocyte cornified envelope, J Biol Chem, 17, 2021, 10.1074/jbc.272.3.2021

Sugarman, 2003, The objective severity assessment of atopic dermatitis score: An objective measure using peermeability barrier function and stratum corneum hydration with computer-assisted estimates for extent of disease, Arch Dermatol, 139, 1417, 10.1001/archderm.139.11.1417

Suzuki, 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, 10.1111/j.1365-2133.1996.tb16230.x

Suzuki, 1993, Detection and characterization of endogenous protease associated with desquamation of stratum corneum, Arch Dermatol Res, 285, 372, 10.1007/BF00371839

Swartzendruber, 1987, Evidence that the corneocyte has a chemically bound lipid envelope, J Invest Dermatol, 88, 709, 10.1111/1523-1747.ep12470383

Takahashi, 1996, Filaggrin linker segment peptide and cystatin alpha are parts of a complex of the cornified envelope of epidermis, Arch Biochem Biophys, 329, 123, 10.1006/abbi.1996.0199

Thompson, 1986, Isolation, properties, and complete amino acid sequence of human secretory leukocyte protease inhibitor, a potent inhibitor of leukocyte elastase, Proc Natl Acad Sci USA, 83, 6692, 10.1073/pnas.83.18.6692

Thune, 1988, The water barrier function of the skin in relation to the water content of stratum corneum, pH and skin lipids. The effect of alkaline soap and syndet on dry skin in elderly, non-atopic patients, Acta Derm Venereol, 68, 277

Traupe, 1989

Uchida, 2000, Epidermal sphingomyelins are precursors for selected stratum corneum ceramides, J Lipid Res, 41, 2071, 10.1016/S0022-2275(20)32369-5

Vallett, 1996, A direct role for sterol regulatory element binding protein in activation of 3-hydroxy-3-methylglutaryl coenzyme A reductase gene, J Biol Chem, 271, 12247, 10.1074/jbc.271.21.12247

van Hooijdonk, 1997, Demonstration of an Na+/H+ exchanger in mouse keratinocytes measured by the novel pH-sensitive fluorochrome SNARF-calcein, Cell Prolif, 30, 351, 10.1046/j.1365-2184.1997.00098.x

Visscher, 2000, Changes in diapered and nondiapered infant skin over the first month of life, Pediatr Dermatol, 17, 45, 10.1046/j.1525-1470.2000.01711.x

Watkinson, 2001, Water modulation of stratum corneum chymotryptic enzyme activity and desquamation, Arch Dermatol Res, 293, 470, 10.1007/s004030100257

Wertz, 1984, Sphingolipids of the stratum corneum and lamellar granules of fetal rat epidermis, J Invest Dermatol, 83, 193, 10.1111/1523-1747.ep12263553

Wertz, 1989, The role of the corneocyte lipid envelope in cohesion of the stratum corneum, J Invest Dermatol, 93, 169, 10.1111/1523-1747.ep12277394

Wiedow, 1990, Elafin: An elastase-specific inhibitor of human skin. Purification, characterization, and complete amino acid sequence, J Biol Chem, 256, 14791, 10.1016/S0021-9258(18)77182-2

Wiedow, 1993, Antileukoprotease in psoriatic scales, J Invest Dermatol, 101, 305, 10.1111/1523-1747.ep12365430

Williams, 1993, From basket weave to barrier. Unifying concepts for the pathogenesis of the disorders of cornification, Arch Dermatol, 129, 626, 10.1001/archderm.1993.01680260096015

Williams, 2003, Enlightened therapy of the disorders of cornification, Clin Dermatol, 21, 269, 10.1016/S0738-081X(03)00059-2

Wood, 1996, Barrier disruption stimulates interleukin-1 alpha expression and release from a pre-formed pool in murine epidermis, J Invest Dermatol, 106, 397, 10.1111/1523-1747.ep12343392

Wood, 1994, Occlusion lowers cytokine mRNA levels in essential fatty acid-deficient and normal mouse epidermis, but not after acute barrier disruption, J Invest Dermatol, 103, 834, 10.1111/1523-1747.ep12413597

Wood, 1994, Barrier function coordinately regulates epidermal IL-1 and IL-1 receptor antagonist mRNA levels, Exp Dermatol, 3, 56, 10.1111/j.1600-0625.1994.tb00047.x

Wood, 1992, Cutaneous barrier perturbation stimulates cytokine production in the epidermis of mice, J Clin Invest, 90, 482, 10.1172/JCI115884

Wood, 1997, Barrier disruption increases gene expression of cytokines and the 55 kD TNF receptor in murine skin, Exp Dermatol, 6, 98, 10.1111/j.1600-0625.1997.tb00154.x

Ye, 2002, Alterations in cytokine regulation in aged epidermis: Implications for permeability barrier homeostasis and inflammation. I. IL-1 gene family, Exp Dermatol, 11, 209, 10.1034/j.1600-0625.2002.110303.x

Yosipovitch, 1993, Skin surface pH in intertriginous areas in NIDDM patients. Possible correlation to candidal intertrigo, Diabetes Care, 16, 560, 10.2337/diacare.16.4.560

Zaiou, 2002, Cathelicidins, essential gene-encoded mammalian antibiotics, J Mol Med, 80, 549, 10.1007/s00109-002-0350-6

Zaiou, 2003, Antimicrobial and protease inhibitory functions of the human cathelicidin (hCAP18/LL-37) prosequence, J Invest Dermatol, 120, 810, 10.1046/j.1523-1747.2003.12132.x

Zeeuwen, 2003, The human cystatin M/E gene (CST6): Exclusion candidate gene for harlequin ichthyosis, J Invest Dermatol, 121, 65, 10.1046/j.1523-1747.2003.12312.x

Zeeuwen, 2001, Cystatin M/E expression is restricted to differentiated epidermal keratinocytes and sweat glands: A new skin-specific proteinase inhibitor that is a target for cross-linking by transglutaminase, J Invest Dermatol, 116, 693, 10.1046/j.1523-1747.2001.01309.x

Zettersten, 1998, Recessive x-linked ichthyosis: Role of cholesterol-sulfate accumulation in the barrier abnormality, J Invest Dermatol, 111, 784, 10.1046/j.1523-1747.1998.00386.x

Zettersten, 1997, Optimal ratios of topical stratum corneum lipids improve barrier recovery in chronologically aged skin, J Am Acad Dermatol, 37, 403, 10.1016/S0190-9622(97)70140-3