Water relations of tetrapod integument

The vertebrate integument represents an evolutionary compromise between the needs for mechanical protection and those of sensing the environment and regulating the exchange of materials and energy. Fibrous keratins evolved as a means of strengthening the integument while simultaneously providing a structural support for lipids, which comprise the principal barrier to cutaneous water efflux in terrestrial taxa. Whereas lipids are of fundamental importance to water barriers, the efficacy of these barriers depends in many cases on structural features that enhance or maintain the integrity of function. Amphibians are exceptional among tetrapods in having very little keratin and a thin stratum corneum. Thus, effective lipid barriers that are present in some specialized anurans living in xeric habitats are external to the epidermis, whereas lipid barriers of amniotes exist as a lipid-keratin complex within the stratum corneum. Amphibians prevent desiccation of the epidermis and underlying tissues either by evaporating water from a superficial aqueous film, which must be replenished, or by shielding the stratum corneum with superficial lipids. Water barrier function in vertebrates generally appears to be relatively fixed, although various species have 'plasticity' to adjust the barrier effectiveness facultatively. While it is clear that both phenotypic plasticity and genetic adaptation can account for covariation between environment and skin resistance to water efflux, studies of the relative importance of these two phenomena are few. Fundamental mechanisms for adjusting the skin water barrier include changes in barrier thickness, composition and physicochemical properties of cutaneous lipids, and/or geometry of the barrier within the epidermis. While cutaneous lipids have been studied extensively in the contexts of disease and cosmetics, relatively little is known about the processes of permeability barrier ontogenesis related to adaptation and environment. Advances in such knowledge have didactic significance for understanding vertebrate evolution as well as practical application to clinical dermatology.