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

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Barrier Function of the Skin: “La Raison d'Être” of the Epidermis Kathi C. Madison Journal of Investigative Dermatology Volume 121, Issue 2, Pages 231-241 (August 2003) DOI: 10.1046/j.1523-1747.2003.12359.x Copyright © 2003 The Society for Investigative Dermatology, Inc Terms and Conditions

Figure 1 Electron micrograph showing the upper stratum granulosum and SC of an organotypic mouse keratinocyte culture. This figure shows a portion of a micrograph previously published inMadison et al (1988). Arrows, keratohyalin granules; arrowheads, intercellular spaces between closely opposed corneocytes; stars, artifactual separation. The intercellular spaces appear empty in this osmium tetroxide postfixed specimen. Original magnification×6000. Journal of Investigative Dermatology 2003 121, 231-241DOI: (10.1046/j.1523-1747.2003.12359.x) Copyright © 2003 The Society for Investigative Dermatology, Inc Terms and Conditions

Figure 2 Structures of the free ceramides of human SC. Numbers 1 to 8 represent thin layer chromatographic mobility with ceramide 1 being the least polar and ceramide 8 the most polar. Ceramide 9 (EOP) has recently been discovered (Ponec et al, 2003) and has a thin layer chromatographic mobility between that of ceramide 2 and ceramide 3. The letters in parentheses give the ceramide classification as suggested inMotta et al (1993). Journal of Investigative Dermatology 2003 121, 231-241DOI: (10.1046/j.1523-1747.2003.12359.x) Copyright © 2003 The Society for Investigative Dermatology, Inc Terms and Conditions

Figure 3 Electron micrograph showing the stacked and patterned lamellar membrane sheets in a single intercellular space in mouse SC postfixed with ruthenium tetroxide. The cornified envelope of the lower corneocyte is clearly visible (arrows); ICS, intercellular space; K, keratin contents of the corneocytes bordering the intercellular space. Original magnification × 200,000. Journal of Investigative Dermatology 2003 121, 231-241DOI: (10.1046/j.1523-1747.2003.12359.x) Copyright © 2003 The Society for Investigative Dermatology, Inc Terms and Conditions

Figure 4 (a) Left: electron micrograph of a single LG in mouse epidermis. A lower magnification view of the same LG was used in a previously published figure (Madison et al, 1987). Arrows, bounding membrane. Original magnification ×300,000. Right: a schematic diagram of a LG as suggested byLandmann (1986). (b) Electron micrograph of extruded LG contents in the intercellular space at the junction of the granular layer and the SC. G, granular cell; ICS, intercellular space; Arrows, granular cell plasma membrane; open arrow, LG. Original magnification ×125,000. Journal of Investigative Dermatology 2003 121, 231-241DOI: (10.1046/j.1523-1747.2003.12359.x) Copyright © 2003 The Society for Investigative Dermatology, Inc Terms and Conditions

Figure 5 Top: structure of acylglucosylceramide. Bottom: proposed model of acylglucosylceramide function in the flattening and stacking of lipid vesicles to generate the double bilayer structure of the internal LG lamellae. The orientation of acylglucosylceramide is not known and both possibilities are shown. The vesicle and double bilayer disk are shown in cross-section. Arrowheads, linoleate moiety; Glu, glucose. Journal of Investigative Dermatology 2003 121, 231-241DOI: (10.1046/j.1523-1747.2003.12359.x) Copyright © 2003 The Society for Investigative Dermatology, Inc Terms and Conditions

Figure 6 The very long chain ω-hydroxy fatty acid-containing ceramides of mammalian epidermis. LG acylglucosylceramides are the precursors of the acylceramides (ceramide 1 (EOS) is shown) in the SC intercellular lamellae and the ω-hydroxyceramides of the lipid envelope. See text for details. Journal of Investigative Dermatology 2003 121, 231-241DOI: (10.1046/j.1523-1747.2003.12359.x) Copyright © 2003 The Society for Investigative Dermatology, Inc Terms and Conditions

Figure 7 Electron micrograph of mouse SC that has been solvent extracted to remove all of the free lipid. A lucent band (the lipid envelope) remains on the exterior surface of the cornified envelopes of adjacent corneocytes. In solvent-extracted SC the lipid envelopes are tightly opposed and a narrow dark line can be seen where they join (seeWertz et al, 1989 for a more detailed discussion). The dominant component of the lucent band is very long chain ω-hydroxyceramides derived from LG acylglucosylceramides (likely from the LG bounding membrane; see text and Figure 6) and covalently bound to cornified envelope proteins. K, keratin contents of two adjacent corneocytes; arrowheads, cornified envelopes. Ruthenium tetroxide postfixation, original magnification ×125,000. Journal of Investigative Dermatology 2003 121, 231-241DOI: (10.1046/j.1523-1747.2003.12359.x) Copyright © 2003 The Society for Investigative Dermatology, Inc Terms and Conditions

Figure 8 Schematic diagram of the epidermis, including the transformations of lipid structures that accompany epidermal differentiation. Keratin filaments are omitted. Not to scale. Journal of Investigative Dermatology 2003 121, 231-241DOI: (10.1046/j.1523-1747.2003.12359.x) Copyright © 2003 The Society for Investigative Dermatology, Inc Terms and Conditions