Dental structure and tooth attachment modes in the common fangtooth Anoplogaster cornuta (Valenciennes, 1833) (Actinopterygii; Trachichthyiformes; Anoplogastridae).

We studied the structure and attachment modes of the teeth of adult Anoplogaster cornuta using light- and scanning-electron microscopic techniques. All teeth were monocuspid, composed solely of orthodentin, and lacked a covering enameloid cap. Fourteen teeth were present in the oral jaws, with three teeth each on the left and right premaxilla and four teeth each on the left and right dentary. The anteriormost premaxillary and dentary teeth were considerably larger than the more posteriorly located ones. The oral jaw teeth were transparent, non-depressible and firmly ankylosed to their respective dentigerous bone by a largely anosteocytic bone of attachment. No evidence for replacement of the large oral jaw teeth was found in the analyzed adult specimens. The bone of attachment exhibited lower calcium and phosphorus concentrations and a higher Ca/P ratio than the orthodentin. The connection between dentinal tooth shaft and bone of attachment was stabilized by a collar of mineralized collagen fibers. In contrast to the oral jaw teeth, the pharyngeal teeth exhibited a ring-like fibrous attachment to their supporting bones. This mode of attachment provides the teeth with some lateral mobility and allows their depression relative to their supporting bones, which may facilitate intra-pharyngeal prey transport. In contrast, a firm ankylosis was observed in numerous small teeth located on the branchial arches. The function of these teeth is presumably to increase the tightness of the pharyngeal basket and thereby the retention of small prey items in a species living in a habitat with only sparse food supply. Our findings corroborate earlier statements on the tooth attachment modes of the oral jaw teeth of Anoplogaster cornuta, but provide new findings for the attachment modes of pharyngeal teeth in this species.

Notes on cranial ontogeny and delayed metamorphosis in the hynobiid salamander Ranodon sibiricus Kessler, 1866 (Urodela).

The skull of larvae, juveniles and adults of the rare and primitive hynobiid salamander Ranodon sibiricus was re-examined using transparencies and illustrated by new graphics. The earliest larva available for investigations already had the dominant bones. The maxillary, however, was still lacking. Previous descriptions regarding the appearance and growth of bones could be largely confirmed. The vomer, first seen as a relatively small obliquely arranged dentate bar in the 3.8 cm long larva, became larger during ontogeny, but did not change its position remarkably. The vomerine pars dentalis with only a single tooth line was straight in larvae and juveniles, but was slightly curved in adults allowing for distinction of an outer and inner portion. This feature is typical and more pronounced in most other hynobiids. The significance of the vomer and vomerine dentition for systematic and phylogenetic purposes and its changes during metamorphosis are briefly discussed. Two of the specimens examined showed delayed metamorphosis very likely caused by low temperatures. Here the temporal course of transformation was "stretched" and therefore some alterations, e.g. regression of the palatinal portion of the palatopterygoid, were shown more clearly. Continuous growth of some skull elements in these individuals suggested a relative independence from metamorphosis perhaps due to variable thyroid activity and/or independent changes in individual tissue sensitivities. It is suggested that remodelling of the mouth roof could be used for staging urodele ontogeny.

Gender-dependent dimorphic teeth in four species of Mesoamerican plethodontid salamanders (Urodela, Amphibia).

The tooth shapes of premaxillary and maxillary teeth of adults of four Mesoamerican salamander species (Urodela, Plethodontidae) were examined. Using scanning electron microscopy we determined whether the monocuspid teeth that are present only on the premaxillary of sexually mature males belong to the conical, unbladed, monocuspid tooth type found in urodele larvae or whether they represent morphological variations of the typical, bladed, bicuspid teeth of metamorphosed individuals. The teeth of some studied species are, in fact, unbladed and in some cases show one very reduced tip. But none of the studied teeth is per definitionem monocuspid and no tooth shows an enameloid layer between dentine shaft and enamel cap, which is always present in the teeth of early urodele larvae. The "monocuspid" teeth of adult males of Mesoamerican plethodontid salamanders have to be considered a morphological variation of the metamorphosed, bicuspid tooth type typical for metamorphosed urodeles.

The dentition of European Speleomantes spp. (Urodela, Plethodontidae) with special regard to sexual dimorphism.

Dentigerous bones and dentition of juveniles and adults of several European Speleomantes-species were investigated using scanning electron microscopy. In the juveniles examined, all dentigerous bones bore bicuspid teeth. The adult males showed the well-known dentitional sexual dimorphism, i.e. monocuspid largely conical, elongated and strongly curved premaxillary teeth. However, teeth of the anterior portion of the maxillae and some teeth of the dentary were also clearly monocuspid; the remaining teeth had differently shaped apices considered to be different grades of bicuspidity. Vomerine teeth were exclusively bicuspid. Monocuspid, but smaller teeth were also found on the premaxilla of an adult female. The conical monocuspid tooth in adults is a modified metamorphosed tooth. Monocuspids, at least on the premaxillae, may be constantly present in males and development of monocuspids probably depends on differential sensitivities of the tooth producing tissue for androgen stimuli.

Separated dental laminae are present in the upper jaw of Mesoamerican lungless salamanders (Amphibia, Plethodontidae).

Plethodontid salamander species from Northern America exhibit a single, unseparated dental lamina in the upper jaw, which consistently provides teeth for the unpaired premaxillary and the two maxillary bones. During the distinct mating season, adult males of these species bear long, monocuspid, unbladed teeth in the entire upper jaw, which are used during courtship. However, plethodontid salamander species from Southern Central America studied so far show either three separated dental laminae or a single dental lamina with three distinguishable tooth-producing parts connected by non-producing parts. These species reproduce aseasonally and sexually mature males permanently bear long, monocuspid, unbladed teeth on the premaxillary only. As the formation of these long, monocuspid teeth depends on the presence of androgen-receptors in the tooth-forming tissues, the morphological separation of the dental lamina in plethodontid salamanders from Southern Central America is very likely to be a prerequisite for a region-specific expression of androgen-receptors in the tooth-forming tissues controlling the differentiated shape formation of teeth in these species. Seven species of plethodontid salamanders from Costa Rica and Panama were examined and, in all, a tendency to exhibit three separate dental lamina in the upper jaw was found, although the morphological separation is definitive only in Bolitoglossa colonnea, B. lignicolor, B. pesrubra, B. schizodactyla and B. striatula. B. dofleini and B. marmorea show a continuous dental lamina with non-producing parts at the transitional sites between the parts associated with the premaxillary and the maxillary bones.

Targeted expression of androgen receptors in tooth-forming tissues of a neotropical salamander (Bolitoglossa schizodactyla) enables region-specific formation of dimorphic types of teeth in the male.

Bicuspid teeth are a morphological characteristic in adult metamorphosed urodeles. However, during the mating season the males of many species of plethodontid salamanders (Urodela, Plethodontidae) from northern America bear large monocuspid teeth on their jaws. Since androgen levels are elevated during the mating season, it is likely that the formation of monocuspid teeth is controlled by androgens. However, many plethodontid salamanders from southern Central America live in habitats which lack significant seasonal changes and therefore are no seasonal breeders. These males constantly bear monocuspid teeth, however their presence is restricted to the premaxillary bone, which forms the most frontal element of the upper jaw. Teeth of all other dentigerous bones are bicuspid. This study is based on an immunohistochemical localisation of androgen receptors and reveals a differential expression of androgen receptors in the three separate dental laminae of the upper jaw of Bolitoglossa schizodactyla (Urodela, Plethodontidae) from western Panama. Our findings suggest that the restriction of androgen receptor expression to the premaxillary dental lamina of the male and its absence in all other tooth-forming tissues leads to the differences in tooth morphogenesis. In conclusion, a differential and cell-specific expression of androgen receptors in urodeles-similar to the situation in mammalian species-allows a selective response of differentiating tissues to the hormonal status of the animal.

Histological studies of bone formation during pedicle restoration and early antler regeneration in roe deer and fallow deer.

The purpose of the present study was to examine the process of bone formation in the regenerating cranial appendages of roe deer (Capreolus capreolus) and fallow deer (Dama dama) during the early postcasting period. After the antlers are cast, osteoclastic and osteoblastic activities lead to a smoothing of the pedicle's separation surface, a strengthening of the pedicle bone, and a partial restoration of the distal pedicle portion that was lost along with the cast antler. Initially, bone formation occurs by intramembranous ossification, but early during the regeneration process cartilage is formed at the tips of the cranial appendages, and is subsequently replaced by bone in a process of endochodral ossification. Shortly after the antlers are cast, the cambium layer of the periosteum in the distal pedicle is markedly enlarged, which suggests that the periosteum serves as a cell source for the bone-forming tissue covering the exposed pedicle bone. The histological findings of our study are consistent with the view that the bony component of the regenerating cranial appendages of deer is largely derived from the pedicle periosteum. Based on findings in other bone systems, we speculate that stem cells that can undergo both osteogenic and chondrogenic differentiation are present in the pedicle periosteum. The early onset of chondrogenesis in the regeneration process is regarded as an adaptation to the necessity of producing a huge volume of bone within a short period. This parallels the situation in other cases of chondrogenesis in membrane bones.

Tooth development in Ambystoma mexicanum: phosphatase activities, calcium accumulation and cell proliferation in the tooth-forming tissues.

Prerequisites of tooth formation, cell proliferation in the tooth-forming tissues, calcium accumulation and the enzymatic activities of alkaline (ALP) and acid phosphatases (ACP) were investigated by immunohistochemical and histochemical methods in various developmental stages of the Mexican Axolotl, Ambystoma mexicanum. During the growth of replacement teeth, the tooth-forming tissues continually recruit cells from the surrounding regions. The basal layer of the oral epithelium, the dental lamina and sometimes even the outer enamel epithelium provide cells for the differentiated inner enamel epithelium, in which the active ameloblasts are localized. The differentiating odontoblasts are derived from proliferating cells situated basally to the replacement teeth in the mesenchymal tissue. When differentiation has started and the cells have become functional, proliferative activity can no longer be observed. Calcium is accumulated close to the site of mineralization in the inner enamel epithelium and in the odontoblasts as it is in mammals, elasmobranchii and teleostei. The activities of ACP and ALP related to the mineralization of the replacement teeth are separated spatially and not sequentially as they are in mammals. However, the results indicate a similar function of these enzymatic components in relation to tooth formation and maturation of mineral deposition. Most of the substantial processes related to tooth formation reported from other vertebrates occur in a manner similar to that in Ambystoma mexicanum, but there also seem to be basic mechanisms present that are realised in a unique way in this urodele.

The skull structure of six species of Mesoamerican plethodontid salamanders (Amphibia, Urodela).

The skulls of six species of plethodontid salamanders of the genus Bolitoglossa from Costa Rica and Panama, Bolitoglossa colonnea, B. dofleini, B. lignicolor, B. marmorea, B. schizodactyla and B. striatula are comparatively analysed. All species are terrestrial or slightly arboreal and show no life-mode-dependent skull characteristics. Heads cleared by transparent preparation and stained in toto were used for examination of the skull structure and paraffin sections of the gonads were prepared to confirm sexual maturity of each individual. Focussing on the size of the premaxillary pars dentalis, the degree of fusion of the processus dorsales praemaxillares and the presence or absence of the paired prefrontals we state that B. dofleini and B. striatula possess more ancestral characteristics, B. marmorea is situated in an intermediate position and B. colonnea, B. lignicolor and B. schizodactyla show more advanced characteristics.

Molting in the Djungarian hamster (Phodopus sungorus Pallas): seasonal or continuous process?

After transfer into a short daylight regimen, the brownish summer pelage of the Djungarian hamster (Phodopus sungorus) changes into the whitish winter phenotype. Although changes in serum prolactin levels are identified as the initiating hormonal signal, morphological data about molting in that species are sparse. The aim of this study was to characterize in detail the summer and winter pelage of the Djungarian hamster and to analyze the alterations in the skin and pelage induced by photoperiodic changes. The main difference between summer and winter hair types is the pattern of pigmentation. In contrast to other mammalian species showing seasonal changes, the winter coat of the Djungarian hamster is not characterized by an increase in hair density. Molting patches were observed at all times, even in the winter coat, showing that the light regimen does not control the process of molting itself but the pattern of pigmentation and eventually the loss of hair during the single molting wave.

The epithelia of the protrusible tongue of Eurycea longicauda guttolineata (Hoolbrook 1838) (Urodela: Plethodontidae).

In this study the lingual and sublingual glands, the lingual stem and the epithelial surface of the protrusible secondary tongue were investigated by light, scanning and transmission electron microscopy. The quality of the secretions of the epithelia was characterized histochemically. The lingual epithelium is formed by superficial (pavement) and goblet cells and at the margin of the tongue pad are also regions covered by ciliated cells. On the dorsal part of the tongue there are goblet cells of type A with mainly acidic secretions and of type B containing neutral secretions. Most of the goblet cells on the ventral side of the tongue (hypoglottis) show a strong alcian blue/PAS positive reaction (type I) and some produce neutral secretions (type II). The glandular cells of the lingual gland react positively to alcian blue and PAS in the apical region of the gland. In contrast there is only alcian blue-positive staining in the basal part of the gland. The size and complexity of the inclusion bodies of the secretory granules increase in a basal direction. In addition, there are ciliated cells in the glandular epithelium. Although the epithelium of the lingual stem is thin, it is double-layered. The cell types observed in this region are identical to those of the ventral part of the protrusible tongue. At the margin of the sublingual gland are trough-like structures. In the center, tubular parts are observed. The cells of this gland are stain strongly with alcian blue (pH 1.0) mainly in the basal part of the gland. The results of this are compared to the tongue pad and the lingual gland of Salamandra salamandra and Ambystoma mexicanum.

Histochemical and ultrastructural studies of cartilage resorption and acid phosphatase activity during antler growth in fallow deer (Dama dama).

Cartilage resorption in forming primary fallow deer antlers was studied by histochemistry and electron microscopy. A high activity of tartrate-resistant acid phosphatase (TRAP), a histochemical marker of skeletal resorbing cells, was first detected in cells located in the mesenchymal tissue separating the columns of hypertrophic cartilage. No cartilage resorption was observed in this region. Intense TRAP staining occurred in large multinucleated cells (identified as inactive osteoclasts) as well as in smaller cells (regarded as mononuclear osteoclast progenitors). On the basis of these findings it was concluded that this was the region where osteoclasts differentiated from progenitor cells. Further proximally, the mineralized cartilage was eroded by active osteoclasts that were located in Howship's lacunae and exhibited an intense TRAP staining. Electron microscopy showed that the cells identified as inactive osteoclasts lacked a polarized organization. In contrast, the active osteoclasts in the zone of cartilage resorption exhibited a typical polarized organization: the nuclei congregated near the basolateral cell surface, and there was a zone of deep membrane infoldings (ruffled border) surrounded by a clear zone at the apical cell pole adjacent to the resorption surface of the mineralized cartilage. The multinucleated cartilage-resorbing cells of the forming antler thus exhibited the typical histochemical and morphological features of active mammalian osteoclasts. Low levels of TRAP activity were also observed in hypertrophic chondrocytes; however, the specificity and potential significance of this staining remain to be elucidated.

[The significance of theramus palatinus to the development of the vomerine bar inSalamandra salamandra (L.)]. / Die Bedeutung desRamus palatinus für die Vomerspangenbildung beiSalamandra salamandra (L.).

The influence of theramus palatinus (VII.) and the postion of the vomer on the vomerine bar, which develops during metamorphosis, was studied by various amputation procedures. Cutting off the nerve cord does not prevent the development of the vomerine bar, as the nerve soon regenerates, whereas cutting off and displacement as well as the removal of portions of theramus palatinus do prevent development. Removal of portions of the nerve cord during metamorphosis produces only parts of the vomerine bar. Obviously an intact ramus palatinus is necessary for the outgrowth of the novel osseous vomerine bar. The course of this outgrowth is independent of the position of the larval vomer.

[Investigations on the development of the vomerine bar inSalamandra salamandra (L.)]. / Untersuchungen zur Bildung der Vomerspange beiSalamandra salamandra (L.).

The development of the vomerine bar inSalamandra salamandra was studied by amputation and transplantation procedures before and during metamorphosis. The bar of the vomer arises from the connective tissue near the caudal margin of the larval vomer. The larval vomer does not regenerate after amputation. A short caudal part of its dental lamina is sufficient for the outgrowing of the complete dental lamina of the vomerine bar. If the tip of the outgrowing vomerine bar is removed, the bar will remain incomplete, which procedure has no influence on the development of the definitive vomer. The material of the developing vomerine bar transplanted to the subepithelial connective tissue of the skin remains intact, but does not continue to grow. Fragments of the skin transplanted to the region from which the bar of the vomer arises prevent the development of the bar.

[Relations between the osseous palate and its dental laminae inSalamandra salamandra (L.) during metamorphosis]. / Beziehungen zwischen Gaumenknochen und ihren Zahnleisten beiSalamandra salamandra (L.) während der Metamorphose.

The relations between the osseous palate and its dental laminae in larvalSalamandra salamandra (L.) during metamorphosis were examined by various amputation procedures. Removal of the mucosa and the subepithelial tissue in the region where the differentiation of the osseous bar of the vomer ("Vomerspange") takes place, does not prevent outgrowth of the bar. When this tissue material is transplanted into the subepithelial connective tissue of the body, structures similar to the bar are not formed. Amputation of the vomer and adhering connective tissue prevents the development of the bar, whereas removal of the palatinal part of the pterygopalatinum has no influence on its development. The vomerine bar does not therefore represent a "Vomeropalatinalspange" in the sense of Wintrebert. The caudal elongation of the dental lamina of the vomer is correlated with the outgrowth of the vomerine bar.