Historically, the term 'keratin' stood for all of the proteins extracted from skin modifications, such as horns, claws and hooves. Subsequently, it was realized that this keratin is actually a mixture of keratins, keratin filament-associated proteins and other proteins, such as enzymes. Keratins were then defined as certain filament-forming proteins with specific physicochemical properties and extracted from the cornified layer of the epidermis, whereas those filamentforming proteins that were extracted from the living layers of the epidermis were grouped as 'prekeratins' or 'cytokeratins'. Currently, the term 'keratin' covers all intermediate filament-forming proteins with specific physicochemical properties and produced in any vertebrate epithelia. Similarly, the nomenclature of epithelia as cornified, keratinized or non-keratinized is based historically on the notion that only the epidermis of skin modifications such as horns, claws and hooves is cornified, that the non-modified epidermis is a keratinized stratified epithelium, and that all other stratified and non-stratified epithelia are non-keratinized epithelia. At this point in time, the concepts of keratins and of keratinized or cornified epithelia need clarification and revision concerning the structure and function of keratin and keratin filaments in various epithelia of different species, as well as of keratin genes and their modifications, in view of recent research, such as the sequencing of keratin proteins and their genes, cell culture, transfection of epithelial cells, immunohistochemistry and immunoblotting. Recently, new functions of keratins and keratin filaments in cell signaling and intracellular vesicle transport have been discovered. It is currently understood that all stratified epithelia are keratinized and that some of these keratinized stratified epithelia cornify by forming a Stratum corneum . The processes of keratinization and cornification in skin modifications are different especially with respect to the keratins that are produced. Future research in keratins will provide a better understanding of the processes of keratinization and cornification of stratified epithelia, including those of skin modifications, of the adaptability of epithelia in general, of skin diseases, and of the changes in structure and function of epithelia in the course of evolution. This review focuses on keratins and keratin filaments in mammalian tissue but keratins in the tissues of some other vertebrates are also considered.

A detailed redescription of the mechanically interacting structural elements of the lingual apparatus of the domestic chicken, Gallus gallus, revealed the functional and constructional role of organized connective tissue (i.e., ligaments and fasciae) as structural elements that ensure the proper biomechanical interactions among the various structures within the lingual apparatus (e.g., cartilaginous and bony skeletal elements, muscles, salivary glands, epithelial structures). Fasciae, together with extrinsic muscles, also connect the lingual apparatus to the other components of the feeding apparatus, such as the skull, jaw apparatus, and larynx. For example, the hyoid apparatus is attached to the skull by a sheath-like fascia (F. vaginalis), the internal structure of which is described here for the first time. Thus, the hyoid suspension in birds differs fundamentally from that in mammals. This study is the first to examine all biomechanically functioning structural elements that are part of the galliform lingual apparatus in a systematic and comprehensive manner. It also provides a set of novel characters that may be useful for future comparative studies in evolutionary and functional morphology.

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The morphology of cornified structures is notoriously difficult to analyse because of the extreme range of hardness of their component tissues. Hence, a correlative approach using light microscopy, scanning electron microscopy, three-dimensional reconstructions based on x-ray computed tomography data, and graphic modeling was applied to study the morphology of the cornified claw sheath of the domesticated cat as a model for cornified digital end organs. The highly complex architecture of the cornified claw sheath is generated by the living epidermis that is supported by the dermis and distal phalanx. The latter is characterized by an ossified unguicular hood, which overhangs the bony articular base and unguicular process of the distal phalanx and creates an unguicular recess. The dermis covers the complex surface of the bony distal phalanx but also creates special structures, such as a dorsal dermal papilla that points distally and a curved ledge on the medial and lateral sides of the unguicular process. The hard-cornified external coronary horn and proximal cone horn form the root of the cornified claw sheath within the unguicular recess, which is deeper on the dorsal side than on the medial and lateral sides. As a consequence, their rate of horn production is greater dorsally, which contributes to the overall palmo-apical curvature of the cornified claw sheath. The external coronary and proximal cone horn is worn down through normal use as it is pushed apically. The hard-cornified apical cone horn is generated by the living epidermis enveloping the base and free part of the dorsal dermal papilla. It forms nested horn cones that eventually form the core of the hardened tip of the cornified claw. The sides of the cornified claw sheath are formed by the newly described hardcornified blade horn, which originates from the living epidermis located on the slanted face of the curved ledge. As the blade horn is moved apically, it entrains and integrates the hard-cornified parietal horn on its internal side. It is covered by the external coronary and proximal cone horn on its external side. The soft-cornified terminal horn extends distally from the parietal horn and covers the dermal claw bed at the tip of the uniguicular process, thereby filling the space created by the converging apical cone and blade horn. The soft-cornified sole horn fills the space between the cutting edges of blade horn on the palmar side of the cornified claw sheath. The superficial soft-cornified perioplic horn is produced on the internal side of the unguicular pleat, which surrounds the root of the cornified claw sheath. The shedding of apical horn caps is made possible by the appearance of microcracks in the superficial layers of the external coronary and proximal cone horn in the course of deformations of the cornified claw sheath, which is subjected to tensile forces during climbing or prey Correspondence Dr Dominique G. Homberger, Department of Biological Sciences, 202 Life Sciences Building, Louisiana State University, Baton Rouge...

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The integumentary musculature of birds consists of three distinct components. The smooth musculature comprises feather and apterial muscles, which form a continuous musculo-elastic layer within the dermis. The feather muscles, which consistently include at least erectors and depressors, interconnect contour feathers within pterylae (i.e., feather tracts) along gridlines that are oriented diagonally to the longitudinal and transverse axes of the body. The apterial muscles interconnect pterylae by attaching to the contour feathers along their peripheries. The striated musculature is composed of individual subcutaneous muscles, most of which attach to contour feathers along the caudal periphery of pterylae A new integrative functional analysis of the integumentary musculature proposes how apterial muscles stabilize the pterylae and modulate the tension of the musculo-elastic layer, and how subcutaneous muscles provide the initial stimulus for erector muscles being able to ruffle the contour feathers within pterylae. It also shows how the arrangement of the contour feathers and integumentary muscles reflects the stresses and strains that act on the avian skin. These mechanical forces are in effect not only in the adult, especially during flight, but may also be active during feather morphogenesis. The avian integument with its complex structural organization may, therefore, represent an excellent model for analyzing the nature of interactions between the environment and genetic material. The predictions of our model are testable, and our study demonstrates the relevance of integrated analyses of complex organs as mechanically coherent systems for evolutionary and developmental biology.