Barrier Function of the Skin: “La Raison d'Être” of the Epidermis

Journal of Investigative Dermatology - Tập 121 Số 2 - Trang 231-241 - 2003
Kathi C. Madison1
1Marshall Dermatology Research Laboratories, Department of Dermatology, University of Iowa, Roy J. and Lucille A. Carver College of Medicine, Iowa City, Iowa, USA

Tóm tắt

Từ khóa


Tài liệu tham khảo

Abraham, 1985, Linoleate-rich acylglucosylceramides of pig epidermis: Structure determination by proton magnetic resonance, J Lipid Res, 26, 761, 10.1016/S0022-2275(20)34334-0

Abraham, 1987, Stratum corneum lipid liposomes: Calcium-induced transformation into lamellar sheets, J Invest Dermatol, 88, 212, 10.1111/1523-1747.ep12525375

Adams, 1993, Effect of organic solvents on in vitro human skin water barrier function, J Invest Dermatol, 101, 609, 10.1111/1523-1747.ep12366068

Arnold, 1988, Ichthosis congenita type III. Clinical and ultrastructural characteristics and distinction within the heterogeneous ichthyosis congenita group, Arch Dermatol Res, 280, 268, 10.1007/BF00440599

Attar, 1997, Inhibition of retinoid signaling in transgenic mice alters lipid processing and disrupts epidermal barrier function, Mol Endocrinol, 11, 792, 10.1210/mend.11.6.0010

Attenborough, 1980

Berenson, 1951, Studies of diffusion of water through dead human skin: The effect of different environmental states and of chemical alterations of the epidermis, Am J Trop Med, 31, 842, 10.4269/ajtmh.1951.s1-31.842

Blank, 1952, Factors which influence the water content of the stratum corneum, J Invest Dermatol, 18, 433, 10.1038/jid.1952.52

Blank, 1953, Further observations on factors which influence the water content of the stratum corneum, J Invest Dermatol, 45, 249, 10.1038/jid.1965.125

Bouwstra, 1999, Cholesterol sulfate and calcium affect stratum corneum lipid organization over a wide temperature range, J Lipid Res, 40, 2303, 10.1016/S0022-2275(20)32105-2

Bouwstra, 2000, The lipid organization in the skin barrier, Acta Derm Venereol, 208, 23

Bouwstra, 2002, Phase behavior of stratum corneum lipid mixtures based on human ceramides: The role of natural and synthetic ceramide 1, J Invest Dermatol, 118, 606, 10.1046/j.1523-1747.2002.01706.x

Bouwstra, 2003, Structure of the skin barrier and its modulation by vesicular formulations, Prog Lipid Res, 42, 1, 10.1016/S0163-7827(02)00028-0

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

Braverman, 1999, Mutations in the gene encoding 3 beta-hydroxysteroid-delta 8, delta 7-isomerase cause X-linked dominant Conradi–Hunermann syndrome, Nat Genet, 22, 291, 10.1038/10357

Breathnach, 1973, Freeze-fracture replication of cells in stratum corneum of human epidermis, J Anat, 114, 65

Cabral, 2001, SPRR4, a novel cornified envelope precursor: UV-dependent epidermal expression and selective incorporation into fragile envelopes, J Cell Sci, 114, 3837, 10.1242/jcs.114.21.3837

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

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

Chilcott, 2002, Transepidermal water loss does not correlate with skin barrier function in vitro, J Invest Dermatol, 118, 871, 10.1046/j.1523-1747.2002.01760.x

De Laurenzi, 1996, Sjögren–Larsson syndrome is caused by mutations in the fatty aldehyde dehydrogenase gene, Nat Genet, 12, 52, 10.1038/ng0196-52

Denda, 1998, Low humidity stimulates epidermal DNA synthesis and amplifies the hyperproliferative response to barrier disruption: Implication for seasonal exacerbations of inflammatory dermatoses, J Invest Dermatol, 111, 873, 10.1046/j.1523-1747.1998.00364.x

Djian, 2000, Targeted ablation of the murine involucrin gene, J Cell Biol, 151, 381, 10.1083/jcb.151.2.381

Doering, 1999, Sphingolipid activator proteins are required for epidermal permeability barrier formation, J Biol Chem, 274, 11045, 10.1074/jbc.274.16.11038

Doering, 1999, Accumulation of protein-bound epidermal glucosylceramides in beta-glucocerebrosidase deficient type 2 Gaucher mice, FEBS Lett, 447, 167, 10.1016/S0014-5793(99)00274-4

Doering, 2002, Sphingolipid metabolism during epidermal barrier development in mice, J Lipid Res, 43, 1727, 10.1194/jlr.M200208-JLR200

Downing, 1992, Lipid and protein structures in the permeability barrier of mammalian epidermis, J Lipid Res, 33, 301, 10.1016/S0022-2275(20)41520-2

Elias, 1975, The permeability barrier in mammalian epidermis, J Cell Biol, 65, 180, 10.1083/jcb.65.1.180

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, 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, Permeability barrier homeostasis. The role of lipid processing, 59

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

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, 2000, The environmental interface. Regulation of permeability barrier homeostasis, 45

Fluhr, 1999, Glycerol accelerates recovery of barrier function in vivo, Acta Derm Venereol, 79, 418, 10.1080/000155599750009825

Forslind, 1994, A domain mosaic model of the skin barrier, Acta Derm Venereol, 74, 1, 10.2340/00015555741214

Freinkel, 1985, Lipid composition and acid hydrolase content of lamellar granules of fetal rat epidermis, J Invest Dermatol, 85, 295, 10.1111/1523-1747.ep12276831

Furuse, 2002, Claudin-based tight junctions are crucial for the mammalian permeability barrier: A lesson from claudin-1-deficient mice, J Cell Biol, 156, 1099, 10.1083/jcb.200110122

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

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

Grange, 2000, CHILD syndrome caused by deficiency of 3 beta-hydroxysteroid-delta 8, delta 7-isomerase, Am J Med Genet, 90, 328, 10.1002/(SICI)1096-8628(20000214)90:4<328::AID-AJMG13>3.0.CO;2-F

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

Hamanaka, 2002, Human epidermal glucosylceramides are major precursors of stratum corneum ceramides, J Invest Dermatol, 119, 416, 10.1046/j.1523-1747.2002.01836.x

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

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

Hansson, 2002, Epidermal overexpression of stratum corneum chymotryptic enzyme in mice: A model for chronic itchy dermatitis, J Invest Dermatol, 118, 444, 10.1046/j.0022-202x.2001.01684.x

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

Harding, 2000, Effects of natural moisturizing factor and lactic acid isomers on skin function, 229

Hart, 1999, Mutations of the cathepsin C gene are responsible for Papillon–Lefevre syndrome, J Med Genet, 36, 881, 10.1136/jmg.36.12.881

Hedberg, 1988, The time course of lipid biosynthesis in pig epidermis, J Invest Dermatol, 91, 169, 10.1111/1523-1747.ep12464438

Hirschberg, 1998, Kinetic analysis of secretory membrane traffic and characterization of Golgi to plasma membrane transport intermediates in living cells, J Cell Biol, 143, 1485, 10.1083/jcb.143.6.1485

Holleran, 1994, Consequences of beta-glucocerebrosidase deficiency in epidermis. Ultrastructure and permeability barrier alterations in Gaucher disease, J Clin Invest, 93, 1756, 10.1172/JCI117160

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

Imokawa, 1991, Stratum corneum lipids serve as a bound-water modulator, J Invest Dermatol, 96, 845, 10.1111/1523-1747.ep12474562

Jain, 2000, Influence of antenatal steroids and sex on maturation of the epidermal barrier in the preterm infant, Arch Dis Child Fetal Neonatal Ed, 83, F112, 10.1136/fn.83.2.F112

Jansen, 1997, Refsum disease is caused by mutations in the phytanoyl-CoA hydroxylase gene, Nat Genet, 17, 190, 10.1038/ng1097-190

Jarnik, 2002, Quasi-normal cornified cell envelopes in loricrin knockout mice imply the existence of a loricrin backup system, J Invest Dermatol, 118, 102, 10.1046/j.0022-202x.2001.01661.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

Jetten, 1992, Increased cholesterol sulfate and cholesterol sulfotransferase activity in relation to the multi-step process of differentiation in human epidermal keratinocytes, J Invest Dermatol, 92, 203, 10.1111/1523-1747.ep12276731

Jobard, 2002, Lipoxygenase-3 (ALOXE3) and 12 (R)-lipoxygenase (ALOX12B) are mutated in non-bullous congenital ichthyosiform erythroderma (NCIE) linked to chromosome 17p13.1, Hum Mol Genet, 11, 107, 10.1093/hmg/11.1.107

Kajava, 2000, Alpha-helical solenoid model for the human involucrin, FEBS Lett, 473, 127, 10.1016/S0014-5793(00)01502-7

Kali, 1998, Development of skin barrier function in premature infants, J Invest Dermatol, 111, 320, 10.1046/j.1523-1747.1998.00289.x

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

Koch, 2000, Lessons from loricrin-deficient mice: Compensatory mechanisms maintaining skin barrier function in the absence of a major cornified envelope protein, J Cell Biol, 151, 389, 10.1083/jcb.151.2.389

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

Konig, 2000, Mutations in the NSDHL gene, encoding a 3beta-hydroxysteroid dehydrogenase, cause CHILD syndrome, Am J Med Genet, 90, 339, 10.1002/(SICI)1096-8628(20000214)90:4<339::AID-AJMG15>3.0.CO;2-5

Kuempel, 1998, In vitro reconstitution of stratum corneum lipid lamellae, Biochim Biophys Acta, 1372, 135, 10.1016/S0005-2736(98)00053-4

Landmann, 1986, Epidermal permeability barrier: Transformation of lamellar granule disks into intercellular sheets by a membrane fusion process, J Invest Dermatol, 87, 202, 10.1111/1523-1747.ep12695343

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

Landmann, 1984, Acylglucosylceramide causes flattening and stacking of liposomes: An analogy for assembly of the epidermal permeability barrier, Biochim Biophys Acta, 778, 412, 10.1016/0005-2736(84)90388-2

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

Lazo, 1999, A mixture of alpha-helical and 310-helical conformations for involucrin in the human epidermal corneocyte envelope provides a scaffold for the attachment of both lipids and proteins, J Biol Chem, 274, 37340, 10.1074/jbc.274.52.37340

Le Fur, 2001, Analysis of circadian and ultradian rhythms of skin surface properties of face and forearm of healthy women, J Invest Dermatol, 117, 718, 10.1046/j.0022-202x.2001.01433.x

Lee, 1998, Iontophoresis itself on hairless mouse skin induces the loss of the epidermal calcium gradient without skin barrier impairment, J Invest Dermatol, 111, 39, 10.1046/j.1523-1747.1998.00226.x

Lefevre, 2001, Mutations in CGI-58, the gene encoding a new protein of the esterase/lipase/thioestserase subfamily, in Chanarin–Dorfman syndrome, Am J Hum Genet, 69, 1002, 10.1086/324121

Loden, 2000, Skin-identical lipids versus petrolatum in the treatment of tape-stripped and detergent-perturbed human skin, Acta Derm Venereol, 80, 412, 10.1080/000155500300012774

Long, 1985, Human stratum corneum polar lipids and desquamation, Arch Dermatol Res, 277, 284, 10.1007/BF00509081

Macheleidt, 2002, Deficiency of epidermal protein-bound omega-hydroxyceramides in atopic dermatitis, J Invest Dermatol, 119, 166, 10.1046/j.1523-1747.2002.01833.x

Madison, 1996, Ceramides are transported through the Golgi apparatus in keratinocytes in vitro, J Invest Dermatol, 106, 1030, 10.1111/1523-1747.ep12338596

Madison, 1987, Presence of intact intercellular lipid lamellae in the upper layers of the stratum corneum, J Invest Dermatol, 88, 714, 10.1111/1523-1747.ep12470386

Madison, 1988, Lamellar granule extrusion and stratum corneum intercellular lamellae in murine keratinocyte cultures, J Invest Dermatol, 90, 110, 10.1111/1523-1747.ep12462065

Madison, 1989, Murine keratinocyte cultures grown at the air/medium interface synthesize stratum corneum lipids and “recycle” linoleate during differentiation, J Invest Dermatol, 93, 10, 10.1111/1523-1747.ep12277335

Madison, 1990, Sphingolipid metabolism in organotypic mouse keratinocyte cultures, J Invest Dermatol, 96, 657, 10.1111/1523-1747.ep12514333

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

Mancini, 1994, Semipermeable dressings improve epidermal barrier function in premature infants, Pediatr Res, 36, 306, 10.1203/00006450-199409000-00007

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

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

Marekov, 1998, Ceramides are bound to structural proteins of the human foreskin epidermal cornified cell envelope, J Biol Chem, 273, 17760, 10.1074/jbc.273.28.17763

Marshall, 2001, Differentially expressed late constituents of the epidermal cornified envelope, Proc Natl Acad Sci USA, 98, 13031, 10.1073/pnas.231489198

Matoltsy, 1968, Studies of the epidermal water barrier. Part II. Investigation of the chemical nature of the water barrier, J Invest Dermatol, 50, 19, 10.1038/jid.1968.3

McCallion, 1995, In vivo evaluation of the effects of moisturisers on transepidermal water loss using factorial designs, Int J Pharmaceutics, 113, 247, 10.1016/0378-5173(94)00207-L

Melton, 1987, Effects of essential fatty acid deficiency on O-acylsphingolipids and transepidermal water loss in young pigs, Biochim Biophys Acta, 921, 191, 10.1016/0005-2760(87)90018-X

Menon, 1986, Cytochemical and biochemical localization of lipase and sphingomyelinase activity in mammalian epidermis, J Invest Dermatol, 86, 591, 10.1111/1523-1747.ep12355263

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

Milner, 1992, Abnormal lamellar granules in harlequin ichthyosis, J Invest Dermatol, 99, 824, 10.1111/1523-1747.ep12614791

Mironov, 1998, Constitutive transport between the trans-Golgi network and the plasma membrane according to the maturation model. A hypothesis, FEBS Lett, 440, 99, 10.1016/S0014-5793(98)01439-2

Morita, 1998, Subcellular distribution of tight junction-associated proteins, occludin, ZO-1 and ZO-2, in rodent skin and its developmental changes, J Invest Dermatol, 110, 862, 10.1046/j.1523-1747.1998.00209.x

Morrison, 1992, ServoMed evaporimeter: Precautions when evaluating the effect of skin care products on barrier function, J Soc Cosmet Chem, 43, 161

Motta, 1993, Ceramide composition of the psoriatic scale, Biochim Biophys Acta, 1182, 145

Motta, 1994, Abnormality of water barrier function in psoriasis: Role of ceramide fractions, Arch Dermatol, 130, 452, 10.1001/archderm.1994.01690040056007

Nemes, 2000, Cholesterol 3-sulfate interferes with cornified envelope assembly by diverting transglutaminase 1 activity from the formation of cross-links and esters to the hydrolysis of glutamine, J Biol Chem, 275, 2636, 10.1074/jbc.275.4.2636

Nemes, 1999, A novel function for transglutaminase 1. Attachment of long-chain omega-hydroxyceramides to involucrin by ester bond formation, Proc Natl Acad Sci USA, 96, 8402, 10.1073/pnas.96.15.8402

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

Onken, 1963, The water barrier in human epidermis, Arch Dermatol, 87, 584, 10.1001/archderm.1963.01590170042007

Orth, 2000, Glycerine: A natural ingredient for moisturizing skin, 213

Pilgram, 2001, Aberrant lipid organization in stratum corneum of patients with atopic dermatitis and lamellar ichthyosis, J Invest Dermatol, 117, 710, 10.1046/j.0022-202x.2001.01455.x

Ponec, 1997, The formation of competent barrier lipids in reconstructed human epidermis requires the presence of vitamin C, J Invest Dermatol, 109, 348, 10.1111/1523-1747.ep12336024

Ponec, 2003, New acylceramide in native and reconstructed epidermis, J Invest Dermatol, 120, 581, 10.1046/j.1523-1747.2003.12103.x

Rigg, 1990, Shed snake skin and hairless mouse skin as model membranes for human skin during permeation studies, J Invest Dermatol, 94, 235, 10.1111/1523-1747.ep12874561

Robson, 1994, 6-Hydroxy-4-sphingenine in human epidermal ceramides, J Lipid Res, 35, 2060, 10.1016/S0022-2275(20)39952-1

Sando, 1996, Induction of ceramide glucosyltransferase in cultured keratinocytes: Correlation with culture differentiation, J Biol Chem, 271, 22044, 10.1074/jbc.271.36.22044

Sato, 1998, Cholesterol sulfate inhibits proteases that are involved in desquamation of stratum corneum, J Invest Dermatol, 111, 189, 10.1046/j.1523-1747.1998.00244.x

Scheuplein, 1970, Effects of surfactants and solvents on the permeability of epidermis, J Soc Cosmet Chem, 21, 853

Schmuth, 2000, Permeability barrier disorder in Niemann-Pick disease: Sphingomyelin-ceramide processing required for normal permeability barrier homeostasis, J Invest Dermatol, 115, 459, 10.1046/j.1523-1747.2000.00081.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

Schmuth, 2002, Role of peroxisome proliferator-activated receptor alpha in epidermal development in utero, J Invest Dermatol, 119, 1298, 10.1046/j.1523-1747.2002.19605.x

Schreiner, 2000, Barrier characteristics of different human skin types investigated with X-ray diffraction, lipid analysis, and electron microscopy imaging, J Invest Dermatol, 114, 654, 10.1046/j.1523-1747.2000.00941.x

Segre, 1999, Klf4 is a transcription factor required for establishing the barrier function of skin, Nature Genet, 22, 356, 10.1038/11926

Shapiro, 1978, X-linked ichthyosis due to steroid sulfatase deficiency, Lancet, i, 70, 10.1016/S0140-6736(78)90005-3

Sidransky, 1992, Gaucher disease in the neonate. A distinct Gaucher phenotype is analogous to a mouse model created by targeted disruption of the beta-glucocerebrosidase gene, Pediatr Res, 32, 494, 10.1203/00006450-199210000-00023

Siegfried, 1998, Neonatal skin and skin care, Dermatol Clin, 16, 437, 10.1016/S0733-8635(05)70244-8

Silverman, 1989, Effects of occlusive and semiocclusive dressings on the return of barrier function to transepidermal water loss in standardized human wounds, J Am Acad Dermatol, 20, 755, 10.1016/S0190-9622(89)70086-4

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

Sondell, 1995, Evidence that stratum corneum chymotryptic enzyme is transported to the stratum corneum extracellular space via lamellar bodies, J Invest Dermatol, 104, 819, 10.1111/1523-1747.ep12607007

Stachowitz, 2002, Permeability barrier disruption increases the level of serine palmitoyltransferase in human epidermis, J Invest Dermatol, 119, 1048, 10.1046/j.1523-1747.2002.19524.x

Steinert, 2000, The complexity and redundancy of epithelial barrier function, J Cell Biol, 151, F5, 10.1083/jcb.151.2.F5

Stewart, 1999, A new 6-hydroxy-4-sphingenine-containing ceramide in human skin, J Lipid Res, 40, 1434, 10.1016/S0022-2275(20)33385-X

Stewart, 2001, The omega-hydroxyceramides of pig epidermis are attached to corneocytes solely through omega-hydroxyl groups, J Lipid Res, 42, 1105, 10.1016/S0022-2275(20)31600-X

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

Swartzendruber, 1989, Molecular models of the intercellular lipid lamellae in mammalian stratum corneum, J Invest Dermatol, 92, 251, 10.1111/1523-1747.ep12276794

Swartzendruber, 1995, Osmium tetroxide and ruthenium tetroxide are complementary reagents for the preparation of epidermal samples for transmission electron microscopy, J Invest Dermatol, 104, 417, 10.1111/1523-1747.ep12665909

Sweeny, 1970, The role of lipids in the epidermal barrier to water diffusion, J Invest Dermatol, 55, 135

Sybert, 1985, Ichthyosis vulgaris: Identification of a defect in filaggrin synthesis correlated with an absence of keratohyalin granules, J Invest Dermatol, 84, 191, 10.1111/1523-1747.ep12264813

Van de Kerkhof, 1995, Epidermal regeneration and occlusion, Acta Derm Venereol, 75, 6, 10.2340/000155557568

Virolainen, 2001, Ultrastructural features resembling those of harlequin ichthyosis in patients with severe congenital ichthyosiform erythroderma, Br J Dermatol, 145, 480, 10.1046/j.1365-2133.2001.04401.x

Walley, 2001, Gene polymorphism in Netherton and common atopic disease, Nat Genet, 29, 175, 10.1038/ng728

Watanabe, 1998, Up-regulation of glucosylceramide synthase expression and activity during human keratinocyte differentiation, J Biol Chem, 273, 9651, 10.1074/jbc.273.16.9651

Welzel, 1995, Occlusion does not influence the repair of the permeability barrier in human skin, Vol. 23, 180

Welzel, 1996, Skin permeability barrier and occlusion: No delay of repair in irritated human skin, Contact Dermatitis, 35, 163, 10.1111/j.1600-0536.1996.tb02335.x

Wertz, 1996, The nature of the epidermal barrier: Biochemical aspects, Adv Drug Del Rev, 18, 283, 10.1016/0169-409X(95)00077-K

Wertz, 1982, Glycolipids in mammalian epidermis. Structure and function in the water barrier, Science, 217, 1261, 10.1126/science.7112128

Wertz, 1983, Ceramides of pig epidermis: Structure determination, J Lipid Res, 24, 759, 10.1016/S0022-2275(20)37950-5

Wertz, 1987, Covalently bound omega-hydroxylacylsphingosine in the stratum corneum, Biochim Biophys Acta, 917, 108, 10.1016/0005-2760(87)90290-6

Wertz, 1990, Metabolism of linoleic acid in porcine epidermis, J Lipid Res, 31, 1839, 10.1016/S0022-2275(20)42327-2

Wertz, 1987, Essential fatty acids and epidermal integrity, Arch Dermatol, 123, 1381, 10.1001/archderm.1987.01660340149038

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

Williams, 2000, Ichthyosis: Where we have been; Disorders of cornification: Where we are going, Curr Probl Dermatol, 12, 171, 10.1016/S1040-0486(00)90011-5

Williams, 1998, Ontogeny of the epidermal permeability barrier, J Invest Dermatol Symp Proc, 3, 75, 10.1038/jidsymp.1998.18

Winsor, 1944, Differential roles of layers of human epigastric skin on diffusion rate of water, Arch Intern Med, 74, 428, 10.1001/archinte.1944.00210240018004

Yardley, 1981, Lipid composition and metabolism in normal and diseased epidermis, Pharmacol Ther, 13, 357, 10.1016/0163-7258(81)90006-1

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