The effects of androgens on the β-glands of the tokay (Gekko gecko): Modification of an hypothesis

Study of the posterior abdominal epidermis in hypophysectomized/thyroidectomized male and female tokays following surgery, and subsequent androgen therapy, indicates that, contrary to a previous model, all aspects of β-gland differentiation are under direct androgenic control. On the other hand, another epidermal specialization, the digital foot-pad, shows a pattern of histogenesis directly comparable to that of β-glands, but is unaffected by androgens. These data are discussed with respect to the evolution of glandular epidermal specializations in gekkonid lizards and the possible role of androgens in modifying the control of cell differentiation in lizard epidermis.     

Lizard and newt tail regeneration: a quantitative study

Almost perfect fits of the Gompertz equation to the growth in length of tail regenerates in the lizard, Lacerta lepida, and the newt, Notophthalmus viridescens, were obtained. Comparison of certain parameters of the equation with published mitotic index data suggests that the Gompertz equation characterizes each system at least from the time that significant mitotic activity is first observed histologically. An objective method for comparing the regeneration periods of the two species is described and applied. A unified hypothesis derived from consideration of properties of the Gompertz equation successfully accounts for the following phenomena reported, but previously unexplained, in the literature: (1) proximal amputations result in longer regenerates than do distal amputations; (2) proximal amputations elicit greater absolute rates of elongation (in mm/day) than do distal amputations; (3) the percent replaced of the length removed is rather constant, regardless of the absolute length regenerated; and (4) one of the parameters of the Gompertz equation appears to be lognormally distributed in a regenerating population. (See text for references.) A computerized interactive graphical system for normalizing growth equations of individual regenerates and integrating the mathematical model with potential candidates for biological control factors is briefly described.     

Histological changes in the epidermis of the subdigital lamellae of Anolis carolinensis during the shedding cycle

The characteristic anoline climbing organ consists of a number of lamellar scales, on whose outer scale surface are numerous keratinized setae which contact the substrate. These setae are derived from the Oberhautchen of the epidermal generation, and as such are renewed and shed periodically along with the rest of the epidermal material. The histological development of the setae is described, and modifications of the surrounding elements are noted. The relative lengths of the setae and their congregation to form a pad unit poses certain mechanical problems during morphogenesis, simply in terms of accommodation between the functional outer epidermal generation and dermal core of each lamella. Regression of the dermal core and a distal migration of some cells permits accommodation within the lamella for the distal aspect of the Oberhautchen layer, or free margin. Additionally, changes in the gross shape of the lamella occur throughout the sloughing cycle, and a swelling of the cells of the lacunar tissue results in a gap between the stratum corneum of inner and outer epidermal generations. There is a considerable amount of variation in mitotic activity between the germinal layers of opposite sides of the lamella.     

Observations on the histochemistry and ultrastructure of regenerating caudal epidermis of the tuatara Sphenodon punctatus (Sphenodontida,Lepidosauria, Reptilia)

Study of the histology, histochemistry, and fine structure of caudal epidermal regeneration in Sphenodon punctatus through restoration of a scaled form reveals that the processes involved resemble those known in lizards. Following establishment of a wound epithelium (WE), subjacent scale neogenesis involves epidermal downgrowths into the dermis. Although the process is extremely slow, and most new scales do not overlap, their epidermal coverings reestablish epidermal generation (EG) formation. As in lizards, the flat, alpha-keratogenic, WE cells contain lipids as revealed by their affinity for Sudan III. A few mucous cells that store large PAS-positive mucus-like granules also occur in WE. During differentiation of WE cells, among the bundles of 70-nm tonofilaments are many lamellar bodies (LBs) and mucous granules (MGs) that discharge their contents into the cytoplasm and extracellular spaces producing a strongly PAS-positive keratinized tissue. Richness of epidermal lipids coexistent with mucus is a primitive characteristic for amniote vertebrates, probably related to functions as a barrier to cutaneous water loss (CWL). As scale neogenesis begins, beneath the superficial WE appear 3-5 layers of irregularly shaped cells. These contain tonofilament bundles surrounded by small, round keratohyalin-like granules (KHLGs) and a keratinized matrix with beta-keratin packets and a 3-5-nm thick keratin granulation. This mixture of alpha- and beta-keratogenic capacities resembles that seen in the innermost cells of a normal tuatara epidermal generation. As in the latter, but in contrast to both normal and regenerating lizard epidermis, no definable shedding complex with interdigitating clear layer and oberhautchen cells occurs (Alibardi and Maderson, 2003). The tortuous boundaries, and merging beta-keratin packets, identify subjacent keratinizing cells as precursors of the typical stratified, squamous beta-layer seen in long-term regenerated caudal skin wherein the entire vertical sequence of epidermal layers resembles that of normal scales. The sequence of events in caudal epidermal regeneration in S. punctatus resembles that documented for lizards. Observed differences between posttrauma scale neogenesis and scale embryogenesis are responses to functional problems involved in, respectively, restoring, or forming, a barrier to CWL while accommodating rapid somatic growth.     

Ultrastructural contributions to an understanding of the cellular mechanisms involved in lizard skin shedding with comments on the function and evolution of a unique Lepidosaurian phenomenon

Previous reports on the fine structure of lizard epidermis are confirmed and extended by SEM and TEM observations of cell differentiation and the form of shed material from the American anole Anolis carolinensis. Attention is drawn to two issues: 1) the tips of the spinules arising from the mature oberhautchen are markedly curved; this morphology can be seen during differentiation; 2) the median keels of scales from all parts of the body show “naked” oberhautchen cells that lack characteristic spinules, but have a membrane morphology comprising a complex system of serpentine microridges. Maderson's ([1966] J. Morphol. 119:39–50) “zip-fastener” model for the role of the shedding complex formed by the clear layer and oberhautchen is reviewed and extended in the light of recent SEM data. Apparently periodic lepidosaurian sloughing permits somatic growth; understanding how the phenomenon is brought about requires integration of data from the organismic to the molecular level. The diverse forms of integumentary microornamentation (MO) reported in the literature can be understood by considering how the cellular events occurring during the renewal phase prior to shedding relate to the emergence of the form-function complex of the β-layer, which provides physical protection. Issues concerning the evolutionary origin of lepidosaurian skin-shedding are discussed. J. Morphol. 236:1–24, 1998. © 1998 Wiley-Liss, Inc.     

Distribution of keratin and associated proteins in the epidermis of monotreme,marsupial, and placental mammals

The expression of acidic and basic keratins, and of some keratinization marker proteins such as filaggrin, loricrin, involucrin, and trichohyalin, is known for the epidermis of only a few eutherian species. Using light and high-resolution immunocytochemistry, the presence of these proteins has been studied in two monotreme and five marsupial species and compared to that in eutherians. In both monotreme and marsupial epidermis lamellar bodies occur in the upper spinosus and granular layers. Development of the granular layer varies between species and regionally within species. There is great interspecific variation in the size (0.1-3.0 microm) of keratohyalin granules (KHGs) associated with production of orthokeratotic corneous tissues. Those skin regions lacking hairs (platypus web), or showing reduced pelage density (wombat) have, respectively, minute or indiscernible KHGs, associated with patchy, or total, parakeratosis. Ultrastructural analysis shows that monotreme and marsupial KHGs comprise irregular coarse filaments of 25-40 nm that contact keratin filaments. Except for parakeratotic tissues of platypus web, distribution of acidic and basic proteins in monotreme and marsupial epidermis as revealed by anti-keratin antibodies AE1, AE2, and AE3 resembles that of eutherian epidermis. Antibodies against human or rat filaggrins have little or no cross-reactivity with epidermal proteins of other mammals: only sparse areas of wombat and rabbit epidermis show a weak immunofluorescence in transitional cells and in the deepest corneous tissues. Of the available, eutherian-derived antibodies, that against involucrin shows no cross-reactivity with any monotreme and marsupial epidermal tissues and that against trichohyalin cross-reacts only with cells in the inner root sheath and medulla of hairs. These results suggest that if involucrin and trichohyalin are present throughout noneutherian epidermis, they may have species-specific molecular structures. By contrast, eutherian-derived anti-loricrin antibodies show a weak to intense cross-reactivity to KHGs and corneous tissues of both orthokeratotic and parakeratotic epidermis in monotremes and marsupials. High-resolution immunogold analysis of loricrin distribution in immature keratinocytes of platypus parakeratotic web epidermis identifies labeled areas of round or irregular, electron-pale granules within the denser keratohyalin component and keratin network. In the deepest mature tissues, loricrin-like labeling is diffuse throughout the cytoplasm, so that cells lack the preferential distribution of loricrin along the corneous envelope that characterizes mature eutherian keratinocytes. Thus, the irregular distribution of loricrin in platypus parakeratotic tissues more resembles that which has been described for reptilian and avian keratinocytes. These observations on the noneutherian epidermis show that a stratum granulosum is present to different degrees, even discontinuous within one tissue, so that parakeratotic and orthokeratotic areas may alternate: this might imply that parakeratotic monotreme epidermis reflects the primitive pattern of amniote alpha-keratogenesis. Absent from anamniote epidermis and all sauropsid beta-keratogenic tissues, the ubiquitous presence of a loricrin-like protein as a major component of other amniote corneous tissues suggests that this is a primitive feature of amniote alpha-keratogenesis. The apparent lack of specific regionalization of loricin near the plasma membranes of monotreme keratinocytes could be an artifactual result of the immunofluorescence technique employed, or there may be masking of the antigenicity of loricrin-like proteins once they are incorporated into the corneous envelope. Nevertheless, the mechanism of redistribution of such proteins during maturation of monotreme keratinocytes is different from, perhaps more primitive, or less specialized, than that in the epidermis of eutherian mammals.     

Born in a follicle--a historical perspective

A review of major studies of tetrapod skin development since the 1870s illustrates how knowledge of structure and mechanism progressed through phases emphasizing Natural History, morphology, endocrinology, and tissue manipulation prior to the prevailing "molecular era." Each successive phase of investigation, while suffering from its own limitations and constraints, has produced conceptual advances. At various times, different systems in various organisms have been research models of choice for practical and/or technical reasons. Comparative studies of scaled and non-scaled integuments and appendages thereof, e.g., nails, claws, glands, hair, and especially feathers, revealed data that suggested new directions for research programs. Some non-mammalian models still offer unique opportunities for pursuit of specific questions pertinent to studies of hair: arguments between American and British schools concerning feather development that originated in the 1930s remain unresolved and may thus affect interpretation of recent investigations. The current emphasis on the study of diffusible molecules involved in papilla-follicle interactions in hair development and replacement can only be understood in the context of the interwoven history of questions relating sequentially to evolutionary homology, physiological controls of tissue homeostasis, embryonic induction, and, most recently, molecular genetics.     

A histological study of the long-term response to trauma of squamate integument

Scale neogenesis following excision of pieces of skin of various sizes and shapes has been studied in several squamate species. In small wounds, neogenesis does not occur, the wound region contracts, and increased areas of hinge differentiation are seen. In large wounds, the time taken for neogenesis to occur, and the size, shape, distribution, and number of the reformed units, are variable. We confirm previous reports that neogenesis accompanies a renewal phase of the shedding cycle, and suggests that this is because both types of morphogenesis require germinal stability. All types of trauma thus far studied show the initial formation of a hyperplastic wound epithelium with the histological characteristics of a normal α-layer: such tissues are the basis for physiological barrier functions. The mechanical functions of the skin can be fairly adequately restored even by abnormal neogenic scales. The observed variability in the process of scale neogenesis, and the final form of the individual units, are argued to be the result of mechanical factors impinging on the differentiating tissues.     

From frog integument to human skin: dermatological perspectives from frog skin biology

For over a century, frogs have been studied across various scientific fields, including physiology, embryology, neuroscience, (neuro)endocrinology, ecology, genetics, behavioural science, evolution, drug development, and conservation biology. In some cases, frog skin has proven very successful as a research model, for example aiding in the study of ion transport through tight epithelia, where it has served as a model for the vertebrate distal renal tubule and mammalian epithelia. However, it has rarely been considered in comparative studies involving human skin. Yet, despite certain notable adaptations that have enabled frogs to survive in both aquatic and terrestrial environments, frog skin has many features in common with human skin. Here we present a comprehensive overview of frog (and toad) skin ontogeny, anatomy, cytology, neuroendocrinology and immunology, with special attention to its unique adaptations as well as to its similarities with the mammalian integument, including human skin. We hope to provide a valuable reference point and a source of inspiration for both amphibian investigators and mammalian researchers studying the structural and functional properties of the largest organ of the vertebrate body.