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Abstract
The primary function of fish skin is to act as a barrier. It provides protection against
physical damage and assists with the maintenance of homoeostasis by minimising exchange
between the animal and the environment. However in some fish, the skin may play a
more active physiological role. This is particularly true in species that inhabit
specialised environmental niches (e.g. amphibious and air-breathing fish such as the
lungfish), those with physiological characteristics that may subvert the need for
the integument as a barrier (e.g. the osmoconforming hagfish), and/or fish with anatomical
modifications of the epidermis (e.g. reduced epithelial thickness). Using examples
from different fish groups (e.g. hagfishes, elasmobranchs and teleosts), the importance
of fish skin as a transport epithelium for gases, ions, nitrogenous waste products,
and nutrients was reviewed. The role of the skin in larval fish was also examined,
with early life stages often utilising the skin as a surrogate gill, prior to the
development of a functional branchial epithelium.
Fish encounter harsh ionic/osmotic gradients on their aquatic environments, and the mechanisms through which they maintain internal homeostasis are more challenging compared with those of terrestrial vertebrates. Gills are one of the major organs conducting the internal ionic and acid-base regulation, with specialized ionocytes as the major cells carrying out active transport of ions. Exploring the iono/osmoregulatory mechanisms in fish gills, extensive literature proposed several models, with many conflicting or unsolved issues. Recent studies emerged, shedding light on these issues with new opened windows on other aspects, on account of available advanced molecular/cellular physiological approaches and animal models. Respective types of ionocytes and ion transporters, and the relevant regulators for the mechanisms of NaCl secretion, Na(+) uptake/acid secretion/NH(4)(+) excretion, Ca(2+) uptake, and Cl(-) uptake/base secretion, were identified and functionally characterized. These new ideas broadened our understanding of the molecular/cellular mechanisms behind the functional modification/regulation of fish gill ion transport during acute and long-term acclimation to environmental challenges. Moreover, a model for the systematic and local carbohydrate energy supply to gill ionocytes during these acclimation processes was also proposed. These provide powerful platforms to precisely study transport pathways and functional regulation of specific ions, transporters, and ionocytes; however, very few model species were established so far, whereas more efforts are needed in other species.
The idea stated in the title, first clearly expressed by August Krogh, is illustrated by examples from animal biochemistry, physiology, general cell biology, experimental medicine, ethology and botany. General aspects of the concept are discussed.
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.
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