Differentiation of the epidermis during scale formation in embryos of lizard

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Abstract
The formation of the first epidermal generations and in particular of the shedding complex in the developing scales of the lizards Anolis lineatopus and Podarcis muralis was studied by electron microscopy. The initially linear and bilayered epidermis turned into symmetric papillae, which became asymmetric scales. The outer periderm was more electrondense than the following layer, provisionally named 'inner periderm' but probably derived from the basal layer. Coarse thick filaments progressively filled the cytoplasm of the inner periderm or formed reticulate bodies resembling avian peridermal granules. Peridermal cells cornified slightly and could be shed in ovo. From the basal layer various suprabasal epithelial layers were produced. The first layer contained keratohyalin-like granules and was identified as a clear layer. Beneath it a spinulated (Anolis) or serrated (Podarcis) oberhäutchen differentiated. The clear and 'oberhäutchen' layers constitute the first shedding complex. The first epidermal layers that were shed consisted of flaking periderm or periderm together with the clear layer. The differentiation of a mesos layer was under way before hatching, when the epidermal morphology resembled a stage 4-5 of the adult shedding cycle. Fibroblasts under the inner side of the scale made few contacts with the basement membrane and their cytoplasmic elongations were mostly oriented parallel to the dense lamina. Instead, fibroblasts under the basal layer of the outer scale surface (BLOS) made numerous contacts with the basement membrane, suggesting that more dermal-epidermal interactions take place on this side of the scale.