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PubMed1994

A physiological measure of carbonic anhydrase in Müller cells.

Carbonic anhydrase activity was characterized in freshly dissociated Müller cells of the salamander retina. Intracellular pH was monitored using ratio imaging of the indicator dye BCECF as extracellular PCO2 was varied. The extracellular solution was switched rapidly (141 ms rise time) from a HEPES buffered to a CO2-HCO3- buffered solution (both pH 7.4). Introduction of CO2-HCO3- produced a rapid cell acidification. Cell pH dropped from a steady-state pH of 7.02 in HEPES solution to pH 6.81 in CO2-HCO3-. Methazolamide, a carbonic anhydrase inhibitor, dramatically reduced the initial rate of acidification, demonstrating that the acidification was produced by the carbonic anhydrase-catalyzed hydration of CO2. The initial rate of acidification, 52.6 pH units per min (0.88 pH units per s), was reduced approximately 150-fold to 0.36 pH units per min by 10(-3) M methazolamide. Half-maximal inhibition occurred at a methazolamide concentration of 5.6.10(-7) M. The carbonic anhydrase inhibitor acetazolamide (10(-3) M) also greatly reduced the rate of cell acidification. The latency to the onset of carbonic anhydrase inhibition was 660 ms for methazolamide and 7.5 s for acetazolamide. The carbonic anhydrase inhibitor benzolamide (10(-4) M, 4 min exposure), which is poorly membrane permeant, had little effect on the rate of cell acidification, indicating that the site of carbonic anhydrase action was intracellular. The activity of Müller cell carbonic anhydrase may help to buffer extracellular CO2 variations in the retina.

Acetazolamide↗
PubMed1995

Calcium waves in dissociated retinal glial (Müller) cells are evoked by release of calcium from intracellular stores.

Calcium imaging techniques were used to study intracellular free calcium ion regulation in isolated Müller cells in response to changes in extracellular potassium concentration and to caffeine and ryanodine. Müller cells were dissociated from the adult tiger salamander (Ambystoma tigrinum) retina and studied using the calcium indicator Fura-2 and video imaging microscopy techniques. Our results demonstrate that elevation of extracellular potassium in the presence of extracellular calcium evokes an increase in intracellular calcium ([Ca2+]i) throughout the length of the Müller cell. In contrast, in the absence of extracellular calcium, elevation of extracellular potassium can trigger a long latency, wave-like increase in [Ca2+]i that begins in the apical region of the Müller cell and moves toward the endfoot. A similar calcium wave can be evoked in Müller cells when they are exposed to caffeine or ryanodine, agents that cause release of calcium from intracellular stores in many cell types. These data suggest that [Ca2+]i may be altered in Müller cells through an extracellular pathway as well as through a ryanodine-sensitive intracellular release mechanism. The functional consequences of these changes in [Ca2+]i remain to be elucidated.

Animals↗
PubMed2002

Skin flaps inhibit both the current of injury at the amputation surface and regeneration of that limb in newts.

For over two decades, we have been investigating a strong (ca. 20-100 microA/cm2), outwardly directed electric current driven through the limb stump for the first few days following amputation in regenerating salamanders. This current is driven through the stump in a proximal/distal direction by the amiloride-sensitive transcutaneous voltage of the intact skin of the stump. Limb regeneration can be manipulated by several technique that manipulate this physiology, demonstrating that the ionic current is necessary, but not sufficient, for normal regeneration of the amphibian limb. Here, we demonstrate that a full thickness graft of skin covering the forelimb stump of newts strikingly inhibits the regeneration of the limb, and that this procedure is also highly correlated to a suppression of peak outwardly directed stump currents in those animals that fail to regenerate.

Amputation Stumps↗
PubMed2002

Female sperm storage in amphibians.

The three orders of extant amphibians are Gymnophiona, Anura, and Urodela. Although all gymnophionans apparently have internal fertilization and many are viviparous, female sperm storage is unknown. Internal fertilization has convergently evolved in a few anurans, but females of just one species, Ascaphus truei, are known to possess oviductal sperm storage tubules (SSTs). The SSTs of A. truei are similar anatomically to such glands in squamate reptiles. This similarity is convergence due to similar functional adaptations and/or internal design constraints. In salamanders and newts (Urodela), absence of sperm storage in females is the ancestral condition (three families). In the derived condition, sperm storage occurs in cloacal glands called spermathecae, and their possession is a synapomorphy for females in the suborder Salamandroidea (seven families). Salamandroids are the only vertebrates with cloacal sperm storage glands. In this paper, a phenetic analysis of variation in spermathecal characters reveals patterns of convergence in certain spermathecal characters in unrelated taxa that breed in similar habitats. In the family Salamandridae, a role in sperm nutrition for the spermathecal epithelium is questioned, and the widespread occurrence of spermiophagy is related to other reproductive strategies. I propose how the packaging of sperm in structurally different types of spermathecae may influence male paternity.

Adaptation, Physiological↗
PubMed1979

The interaction of nerves of opposite regenerating polarity in fused newt forelimbs.

Previous studies involving nerve interactions and limb regenerative processes were carried out on adult newts after their forelimbs were amputated through the distal radius and ulna and fused end-to-end. On the basis of limb regeneration results at the junction of the fused limbs, it was postulated that regenerating nerves from each limb (i.e., nerves of opposite polarity) would not invade the foreign territory of the contralateral limb if it were already normally innervated. A direct study of this nerve interaction, however, was not made in this earlier study. The present investigation was designed to obtain direct histological and electrophysiological evidence for the interaction of nerves of opposite regenerating polarity in fused newt forelimbs. The primary objective was to determine how the regenerating nerves would interact in the establishment of innervation territories-first, at the fusion zone, which represents the junction of the normal innervation territories of the nerves of each limb; and secondly, half way up one of the limbs, where interaction would occur in a territory normally innervated by only one of the regenerating nerves. The results showed that when nerves of opposite regenerating polarity approached one another at the junction of the fused limbs a discontinuation of axonal growth occurred; no indication of overlap of nerves into foreign territory was seen. When the nerves were allowed to interact within one of the fused limbs, however, an overlap of nerve fibers and a functional "double innervation" of that limb was demonstrated. These results are discussed in terms of possible mechanisms for the establishment of innervation territories in salamander limbs. The question of nerve-muscle reinnervation specificity is also raised.

Animals↗
PubMed1979

Reduction of sodium dependent stump currents disturbs urodele limb regeneration.

We have asked the question whether the natural electric currents which leave urodele limb stumps are in any way needed for their regeneration. As an initial test, we have greatly reduced such currents in the tiger salamander, Ambystoma tigrinum, by applying 0.5 mM amiloride to the stump skin or by immersion of the animals in sodium depleted media. We have also reduced such currents in the red spotted newt, Notophthalmus viridescens, by such immersion. Limb regeneration in half of the amiloride-treated animals was either entirely blocked or grossly deficient, while the others regenerated normally. Limb regeneration in sodium depleted media was consistently inhibited for some weeks but then recovered. These results are consistent with the hypothesis that stump currents are in some way needed for normal regeneration.

Ambystoma↗
PubMed1981

Hormone-induced ovulation in Ambystoma tigrinum: influence of prolactin and thyroxine.

Treatment of hypophysectomized or intact neotenic tiger salamander larvae (Ambystoma tigrinum) with ovine prolactin (PRL) increased sensitivity of the ovary to the ovulation-inducing hormones, human chorionic gonadotropin and luteinizing hormone (LH). Effects of pretreatment in vivo with PRL or thyroxine (T4) on the responsiveness of ovarian fragments in vitro to ovulation-inducing agents, LH, or progesterone (PROG) in the presence or absence of PRL were observed. In November (prior to normal spawning occurring from January through May) in vivo and in vitro treatment with PRL increased the number of oocytes producing polar bodies and the number of eggs ovulated in vitro. In April addition of PRL in vitro increased polar body formation and ovulations in response to LH or PROG, but in vivo pretreatment with PRL had no effect. Pretreatment in vivo with T4 blocked the in vitro enhancement observed with PRL. Thus, endogenous PRL may play a role in the development of ovarian sensitivity to ovulatory hormones at the level of the oocyte, and T4 may inhibit this action.

Ambystoma↗
PubMed1984

Cell cycle and histological effects of reinnervation in denervated forelimb stumps of larval Ambystoma.

Although nerves are a requirement for regeneration of a salamander limb, denervation of an amputated larval Ambystoma forelimb does not permanently prevent the limb from regenerating. When nerves grow back into the distal stump in sufficient numbers, regeneration occurs. In this study we examined histological and cell cycle events in denervated limb stumps of Ambystoma as they became reinnervated and began to regenerate. Prior to reinnervation and regeneration, dedifferentiated cells are present in the distal stump but are rarely synthesizing DNA or undergoing mitosis and are densely packed. About 2 days after the distal tip of the stump becomes reinnervated, the limb tip appears to "expand" as cells become less densely packed, and cell cycle activity begins. Mitotic index (MI) and labeling index (LI) increase slowly compared to MI and LI in control amputated limbs. Evidence for the localization of cycling activity is presented. In reinnervated stumps, the increase in LI precedes the increase in MI, suggesting that most of the cells are initiating cycling from the G1 phase.

Ambystoma↗
PubMed1992

Autocrine and paracrine role of steroids during spermatogenesis: studies in Squalus acanthias and Necturus maculosus.

Due to a cystic mode of spermatogenesis and zonal testes, the spiny dogfish shark (Squalus acanthias) and salamander (Necturus maculosus) are advantageous for studying the relationship between steroidogenesis and germ cell development. Direct analysis of steroidogenic enzymes and steroid receptors in staged tissue samples, together with light and electron microscopic observation, suggests that androgens and estrogens produced by Sertoli cells or Leydig cells immediately before or after spermiation serve as intratesticular regulators of events specific to premeiotic stages of spermatogenesis. These correlative data are consistent with limited observations in mammalian testis and provide a conceptual framework for direct testing of functional interactions between spermatogenesis and steroidogenesis in future studies.

Animals↗
PubMed2007

A Bayesian approach on molecules and behavior: reconsidering phylogenetic and evolutionary patterns of the Salamandridae with emphasis on Triturus newts.

The monophyly of European newts of the genus Triturus within the family Salamandridae has for decades rested on presumably homologous behavioral and morphological characters. Molecular data challenge this hypothesis, but the phylogenetic position of Triturus within the Salamandridae has not yet been convincingly resolved. We addressed this issue and the temporal divergence of Triturus within the Salamandridae with novel Bayesian approaches applied to DNA sequence data from three mitochondrial genes (12S, 16S and cytb). We included 38 salamandrid species comprising all 13 recognized species of Triturus and 16 out of 17 salamandrid genera. A clade comprising all the "Newts" can be separated from the "True Salamanders" and Salamandrina clades. Within the "Newts" well-supported clades are: Tylototriton-Pleurodeles, the "New World Newts" (Notophthalmus-Taricha), and the "Modern Eurasian Newts" (Cynops, Pachytriton, Paramesotriton=together the "Modern Asian Newts", Calotriton, Euproctus, Neurergus and Triturus species). We found that Triturus is a non-monophyletic species assemblage, which includes four groups that are themselves monophyletic: (i) the "Large-Bodied Triturus" (six species), (ii) the "Small-Bodied Triturus" (five species), (iii) T. alpestris and (iv) T. vittatus. We estimated that the last common ancestor of Triturus existed around 64 million years ago (mya) while the root of the Salamandridae dates back to 95 mya. This was estimated using a fossil-based molecular dating approach and an explicit framework to select calibration points that least underestimated their corresponding nodes. Using the molecular phylogeny we mapped the evolution of life history and courtship traits in Triturus and found that several Triturus-specific courtship traits evolved independently.

Animals↗
PubMed2007

Is Palaeospondylus gunni a fossil larval lungfish? Insights from Neoceratodus forsteri development.

The enigmatic Devonian fossil Palaeospondylus gunni was identified as a larval form, metamorphosing into the lungfish Dipterus valenciennesi. Morphological features used to identify P. gunni as a larval lungfish include enlarged cranial ribs, rudimentary limb girdles, and absence of teeth. However, this combination of features does not characterize the extant lungfish Neoceratodus forsteri, even at very young stages, nor early stages of Devonian and younger fossil lungfish. Absence of teeth is problematic because early ontogenetic stages of fossil and living lungfish possess full dentitions including marginal teeth. Also problematic are cranial ribs as a defining character of lungfish, as these also occur in certain actinopterygians. It is argued that Neoceratodus is an obligate neotene (reproductively mature larva), with the implication that metamorphosis was a feature of the ontogeny of early lungfish. Pedomorphic characters have been recognized in Neoceratodus and other post-Devonian lungfish, including large cells and correspondingly large genome size; these latter characters correlate with neoteny in salamanders. Small cells preserved in fossil bone suggest that Devonian lungfish had a smaller genome than post-Devonian lungfish, implying that they were not neotenic. As fossil lungfish cell sizes (and genomes) increased in the late Paleozoic, the diversity of lungfish morphologies decreased, so that taxa like Sagenodus and Conchopoma show morphological similarity to Neoceratodus, marking a point in phylogeny at which metamorphosis was potentially lost. Since ancestral larval characters are retained in neotenic adults, we predict that Devonian larvae should resemble these post-Devonian taxa, a prediction which Palaeospondylus does not fulfill.

Animals↗
PubMed1994

Differential effects of temperature on macrophages of ectothermic vertebrates.

The role of macrophages in vertebrate defense reactions is especially important at low temperatures, which commonly affect representatives of ectothermic species. The aim of this work was to compare the effects of in vivo thermal acclimation and in vitro assay temperatures on peritoneal macrophages (PMs) from fish (carp and goldfish) and amphibians (salamanders and toads). The substratum adherence of PMs was undisturbed over the wide range of temperatures experienced by animals in nature. Dehydrogenases and endocytosis were sensitive to temperature fluctuations in a species-specific manner, more or less dependent on the previous thermal history of animals. The MTT reduction, inhibited by low assay temperatures, was slightly improved in PMs from cold-acclimated amphibians but not fishes. Endocytosis by the amphibian PMs, very efficient over the wide range of assay temperatures, was additionally improved by previous in vivo cold exposure. Contrasting effects of thermal acclimation were most evident in the case of endocytic properties of fish PMs; endocytosis by PMs from cold-acclimated fishes was extraordinarily efficient at a temperature just above zero, which almost completely inhibited cells from fishes adapted to warmth.

Animals↗
PubMed2006

Morphological variations in a tooth family through ontogeny in Pleurodeles waltl (Lissamphibia, Caudata).

Most nonmammalian species replace their teeth continuously (so-called polyphyodonty), which allows morphological and structural modifications to occur during ontogeny. We have chosen Pleurodeles waltl, a salamander easy to rear in the laboratory, as a model species to establish the morphological foundations necessary for future molecular approaches aiming to understand not only molecular processes involved in tooth development and replacement, but also their changes, notably during metamorphosis, that might usefully inform studies of modifications of tooth morphology during evolution. In order to determine when (in which developmental stage) and how (progressively or suddenly) tooth modifications take place during ontogeny, we concentrated our observations on a single tooth family, located at position I, closest to the symphysis on the left lower jaw. We monitored the development and replacement of the six first teeth in a large growth series ranging from 10-day-old embryos (tooth I1) to adult specimens (tooth I6), using light (LM), scanning (SEM), and transmission electron (TEM) microscopy. A timetable of the developmental and functional period is provided for the six teeth, and tooth development is compared in larvae and young adults. In P. waltl the first functional tooth is not replaced when the second generation tooth forms, in contrast to what occurs for the later generation teeth, leading to the presence of two functional teeth in a single position during the first 2 months of life. Larval tooth I1 shows dramatically different features when compared to adult tooth I6: a dividing zone has appeared between the dentin cone and the pedicel; the pulp cavity has enlarged, allowing accommodation of large blood vessels; the odontoblasts are well organized along the dentin surface; tubules have appeared in the dentin; and teeth have become bicuspidate. Most of these modifications take place progressively from one tooth generation to the next, but the change from monocuspid to bicuspid tooth occurs during the tooth I3 to tooth I4 transition at metamorphosis.

Animals↗
PubMed1990

Comparative studies on the vascular organization of carotid labyrinths of anurans and caudates.

The three-dimensional structures of the carotid labyrinth in five species of anurans representing four families (Rana nigromaculata, Rana catesbeiana, Bufo japonicus, Hyla arborea, and Xenopus laevis), and three species of caudates representing three families (Cynops pyrrhogaster, Hynobius nebulosus, Ambystoma mexicanum) were compared using vascular corrosion castings and scanning electron microscopy (SEM). Anuran carotid labyrinths are spherical in shape and are classified into two groups according to the origin of the external and internal carotid arteries. One group, which included Rana, Hyla, and Bufo, is characterized by the presence of a vascular ring at the proximal end and some vascular routes at the distal end of the labyrinth. The external and internal carotid arteries originate from these structures. The other group, which includes only Xenopus, is characterized by the external carotid artery opening directly from the central chamber or the common carotid artery, and by the internal carotid artery originating from within the vascular maze. The vascular maze is most complex in Xenopus, less so in Rana and Bufo, and simplest in Hyla. The carotid labyrinths in Cynops and Hynobius are oblong in shape. The fundamental organization in salamanders is similar to that in anurans. The vascular maze, however, is much simpler than in Hyla. There is no specialized swelling in Ambystoma mexicanum. The present findings suggest that most amphibian carotid labyrinths have the appropriate architecture for controlling vascular tone.

Animals↗
PubMed1998

Variance in male mating success and female choice for persuasive courtship displays.

An important function of the courtship displays of male animals is to stimulate, or 'persuade', females to mate. While many studies of courtship success have shown that females choose stimulating males as partners, fewer have demonstrated that the most persuasive males in a population also enjoy the highest mating success (i.e. mate with the greatest numbers of partners). Such a relationship provides a stronger test of the hypothesis that male displays evolve by sexual selection via female choice. We tested this hypothesis in the laboratory using the North American plethodontid salamander, Desmognathus ocoee, in which females only elicit spermatophore deposition if rendered sexually responsive by male courtship. First, we determined variance among individual males in the numbers of females with which they mated across 35 encounters, in the absence of differences in partner encounter rates and direct intermale interactions. Second, we demonstrated that males with high 'historical' scores of mating success performed significantly higher frequencies of persuasive courtship displays, which provide tactile, chemical and visual stimulation to females. We conclude that sexual selection via female choice favours persuasive displays because they confer high mating success on the males that perform them at the highest frequencies. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗
PubMed1993

Expression of wnt10a in the central nervous system of developing zebrafish.

A polymerase chain reaction-derived clone with sequence homology to partial-length shark and salamander Wnt-10a was used to screen a zebrafish cDNA library, yielding a cDNA encoding a full-length Wnt polypeptide. Transcripts encoding zebrafish wnt10a are first detected during the segmentation period of embryogenesis, at a time when there is extensive development and differentiation of the central nervous system. wnt10a transcripts are confined to differentiating brain regions, with expression first appearing diffusely in the floor of the hindbrain ventricle and proceeding anteriorly to the mid- and forebrain. Comparatively, zebrafish wnt1 transcripts are expressed in a stripe at the future midbrain-hindbrain boundary and at the future forebrain-midbrain border. With the differentiation of the neural tube into distinct regions, wnt1 signals are also detected along the dorsal midline of the midbrain, the mid- and posterior hindbrain, and the spinal cord. These patterns suggest roles for wnt10a in the regionalization of the brain.

Amino Acid Sequence↗
PubMed1994

Developmental regulation of a matrix metalloproteinase during regeneration of axolotl appendages.

Removal of specific extracellular matrix (ECM) components has been implicated in the initiation of salamander limb regeneration. Remodeling of the ECM at the distal stump is necessary for the release of cells that eventually contribute to the blastema. Several matrix metalloproteinases (MMPs) have been well characterized as important to various physiological and pathological processes, such as bone remodeling and tumor invasion. The goal of this study is to identify and characterize MMPs that modulate the ECM during appendage regeneration in the axolotl Ambystoma mexicanum. By analyzing axolotl tissue extracts using gelatin-substrate gels, we have identified a 90-kDa gelatinase/collagenase that is upregulated within hours after limb amputation. This gelatinase shows a dramatic elevation in activity during the dedifferentiation and blastema stages. Its activity declines by the palette stage and returns to its basal level by the digit stages. The increase in activity of the 90-kDa gelatinase in response to amputation is independent of the nerve supply and the wound epithelium but these factors affect its subsequent downregulation. In addition to the blastema, the 90-kDa gelatinase can be detected in the stump at least 4 mm proximal to the regenerate. Similar regulation of the 90-kDa gelatinolytic activity is observed during tail regeneration and flank would healing. We suggest that this 90-kDa gelatinase/collagenase may play a role in the initiation and rapid growth phase of axolotl regeneration and wound healing.

Ambystoma↗
PubMed1995

Electroreceptors and mechanosensory lateral line organs arise from single placodes in axolotls.

The lateral line system in salamanders consists of mechanoreceptive neuromasts and pit organs, distributed in lines on the head and trunk, and electroreceptive ampullary organs located adjacent to the cephalic lines of mechanoreceptors. Although numerous studies have documented that neuromast and pit organs and the cranial nerves that innervate these receptors arise from a dorsolateral series of placodes, there is no agreement concerning the number of these placodes, the specific groups of receptors that arise from them, or the embryonic origin of ampullary organs. A developmental model was recently proposed (Northcutt et al., 1994) in which all these placodes, except for the most posterior one, elongate to form sensory ridges whose central zones initially form neuromast and pit organ primordia and whose lateral zones subsequently form ampullary primordia. To test this model, individual placodes were unilaterally extirpated, or placodes from pigmented wild-type axolotl embryos were homotopically or heterotopically transplanted into albino hosts. Extirpation resulted in the loss of all three receptor classes, and both homotopic and heterotopic transplants produced pigmented receptors of all three classes in albino hosts. The receptors in the heterotopic transplants still formed lines which occasionally retained their normal orientation despite differentiating in an ectopic environment. These experiments demonstrated that, as previously postulated, specific lines of neuromasts and pit organs do arise from each placode, and ampullary organs also arise from many of the same placodes. The distribution of receptors that develop following incomplete extirpation or heterotopic transplantation also indicates that each placode is patterned regarding receptor classes and orientation prior to sensory ridge formation.

Ambystoma↗