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PubMed2002

Two second messengers mediate amino acid responses in olfactory sensory neurons of the salamander, Necturus maculosus.

Odor transduction mediated by the adenylyl cyclase/cAMP pathway has been well studied, but it is still uncertain whether this pathway mediates the transduction of all odors in vertebrates. We isolated olfactory sensory neurons from the salamander Necturus maculosus and used calcium imaging with the indicator dye fura-2 to examine olfactory responses elicited by amino acids. The properties of approximately two-thirds of the odor responses suggested they were mediated by the adenylyl cyclase/cAMP pathway, but one-third of the responses were not mimicked by cAMP analogs nor blocked by inhibition of adenylyl cyclase, suggesting that these odor responses were mediated differently. Responses that were unaffected by inhibition of adenylyl cyclase were blocked by neomycin, an inhibitor of phospholipase C, implying that they were transduced by activation of phospholipase C. Some cells which responded to more than one amino acid appeared to employ both pathways, but each was used to transduce different odors. In addition, many responses that were mediated by the adenylyl cyclase/cAMP pathway were enhanced following inhibition of phospholipase C, suggesting that the phospholipase C pathway has a role not only in odor transduction, but also in the modulation of olfactory responses.

Action Potentials↗
PubMed1995

Separating population structure from population history: a cladistic analysis of the geographical distribution of mitochondrial DNA haplotypes in the tiger salamander, Ambystoma tigrinum.

Nonrandom associations of alleles or haplotypes with geographical location can arise from restricted gene flow, historical events (fragmentation, range expansion, colonization), or any mixture of these factors. In this paper, we show how a nested cladistic analysis of geographical distances can be used to test the null hypothesis of no geographical association of haplotypes, test the hypothesis that significant associations are due to restricted gene flow, and identify patterns of significant association that are due to historical events. In this last case, criteria are given to discriminate among contiguous range expansion, long-distance colonization, and population fragmentation. The ability to make these discriminations depends critically upon an adequate geographical sampling design. These points are illustrated with a worked example: mitochondrial DNA haplotypes in the salamander Ambystoma tigrinum. For this example, prior information exists about restricted gene flow and likely historical events, and the nested cladistic analyses were completely concordant with this prior information. This concordance establishes the plausibility of this nested cladistic approach, but much future work will be necessary to demonstrate robustness and to explore the power and accuracy of this procedure.

Ambystoma↗
PubMed2004

Stabilizing selection on behavior and morphology masks positive selection on the signal in a salamander pheromone signaling complex.

Natural selection maintains the integration and coordination of sets of phenotypic characters that collectively perform a task. In functional complexes in which characters span molecular to behavioral levels of organization, we might then expect similar modes of selection to produce similar patterns in evolutionary divergence at each level. To test this expectation, we diagnosed selection at behavioral, morphological, and molecular levels for courtship pheromone signaling by plethodontid salamanders. At the levels of morphology and behavior tens of millions of years of stasis (stabilizing selection) occur on each side of a transition from vaccination to olfactory delivery modes. As a proxy for the molecular level, we used plethodontid receptivity factor (PRF), a protein that is an active component of the pheromone. We cloned PRF from 12 Plethodon spp. spanning the delivery transition and obtained multiple alleles from each individual surveyed. Analyses of 61 alleles for PRF identified elevated nonsynonymous over synonymous substitution rates along lineages in a molecular phylogeny, and at 8% of sites in the protein, indicating that positive (directional) selection has acted on this vertebrate pheromone gene. Structural models showed PRF is in a family of cytokines characterized by a four-alpha-helix bundle. Positive selection in PRF was associated with receptor binding sites that are under purifying selection in other cytokines of that family. The evolutionary dynamics of the plethodontid pheromone delivery complex consists of stabilizing selection on morphological and behavioral aspects of signal delivery but positive selection on the signal mediated by receptors. Thus, different selection modes prevail at different levels in this reproductive functional complex. Evolutionary studies of integrated sets of characters therefore require separate analyses of selective action at each level.

Amino Acid Sequence↗
PubMed2005

Molecular mechanisms of extensive mitochondrial gene rearrangement in plethodontid salamanders.

Extensive gene rearrangement is reported in the mitochondrial genomes of lungless salamanders (Plethodontidae). In each genome with a novel gene order, there is evidence that the rearrangement was mediated by duplication of part of the mitochondrial genome, including the presence of both pseudogenes and additional, presumably functional, copies of duplicated genes. All rearrangement-mediating duplications include either the origin of light-strand replication and the nearby tRNA genes or the regions flanking the origin of heavy-strand replication. The latter regions comprise nad6, trnE, cob, trnT, an intergenic spacer between trnT and trnP and, in some genomes, trnP, the control region, trnF, rrnS, trnV, rrnL, trnL1, and nad1. In some cases, two copies of duplicated genes, presumptive regulatory regions, and/or sequences with no assignable function have been retained in the genome following the initial duplication; in other genomes, only one of the duplicated copies has been retained. Both tandem and nontandem duplications are present in these genomes, suggesting different duplication mechanisms. In some of these mitochondrial DNAs, up to 25% of the total length is composed of tandem duplications of noncoding sequence that includes putative regulatory regions and/or pseudogenes of tRNAs and protein-coding genes along with the otherwise unassignable sequences. These data indicate that imprecise initiation and termination of replication, slipped-strand mispairing, and intramolecular recombination may all have played a role in generating repeats during the evolutionary history of plethodontid mitochondrial genomes.

Animals↗
PubMed2005

Lineage-specific differences in evolutionary mode in a salamander courtship pheromone.

Functionally equivalent genes may evolve heterogeneously across closely related taxa as a consequence of lineage-specific selective pressures. Such disparate evolutionary modes are especially prevalent in genes that encode postcopulatory reproductive proteins, presumably as a result of sexual selection. We might therefore expect genes that mediate reproduction prior to insemination to evolve in a similar manner. Plethodontid receptivity factor (PRF), a proteinaceous salamander pheromone produced by the male, increases female receptivity during courtship interactions. To test for lineage-specific differences in PRF's evolution, we intensively sampled PRF genes across the eastern Plethodon phylogeny (27 spp.; 34 populations) to compare gene diversification, rates of evolution, modes of selection, and types of amino acid substitution. Our analyses indicate that PRF evolutionary dynamics vary considerably from lineage to lineage. Underlying this heterogeneity, however, are two well-defined transitions in evolutionary mode. The first mode is representative of a typical protein profile, wherein neutral divergence and purifying selection are the dominant features. The second mode is characterized by incessant, cyclical evolution driven by positive selection. In this mode, the positively selected sites are bound by a limited assortment of acceptable amino acids that appear to evolve independently of other sites, resulting in a tremendous number of unique PRF alleles. Several of these selected sites are implicated in receptor binding. These sites are apparently involved in a molecular tango in which the male signal and female receptors coevolve within a confined molecular space. PRF's lineage-specific evolutionary dynamics, in combination with evidence of a molecular tango, highlight the molecular action of sexual selection on a chemical signal that is used during courtship.

Amino Acid Sequence↗
PubMedDate not supplied

Allozyme heterozygosity and growth in the tiger salamander, Ambystoma tigrinum.

Statistically significant, positive correlations were found between individual heterozygosity at seven protein polymorphisms and size in five natural populations of tiger salamanders. When these same populations were sampled later in the same year, the positive associations between heterozygosity and size were no longer evident. A pair mating was conducted in the laboratory, and offspring were distributed randomly to four replicate populations maintained in the laboratory. In two to these four replicates, a significant, positive association between protein heterozygosity and size was observed. The laboratory results suggest that the associations seen in natural populations are due to associations between protein heterozygosity and growth rate early in the larval period.

Ambystoma↗
PubMed1997

Large, rapidly evolving intergenic spacers in the mitochondrial DNA of the salamander family Ambystomatidae (Amphibia: Caudata).

We report the presence, in the mitochondrial DNA (mtDNA) of all of the sexual species of the salamander family Ambystomatidae, of a shared 240-bp intergenic spacer between tRNAThr and tRNAPro. We place the intergenic spacer in context by presenting the sequence of 1,746 bp of mtDNA from Ambystoma tigrinum tigrinum, describe the nucleotide composition of the intergenic spacer in all of the species of Ambystomatidae, and compare it to other coding and noncoding regions of Ambystoma and several other vertebrate mtDNAs. The nucleotide substitution rate of the intergenic spacer is approximately three times faster than the substitution rate of the control region, as shown by comparisons among six Ambystoma macrodactylum sequences and eight members of the Ambystoma tigrinum complex. We also found additional inserts within the intergenic spacers of five species that varied from 87-444 bp in length. The presence of the intergenic spacer in all sexual species of Ambystomatidae suggests that it arose at least 20 MYA and has been a stable component of the ambystomatid mtDNA ever since. As such, it represents one of the few examples of a large and persistent intergenic spacer in the mtDNA of any vertebrate clade.

Ambystoma↗
PubMed1997

Phylogenetic relationships of bolitoglossine salamanders: a demonstration of the effects of combining morphological and molecular data sets.

We analyzed sequence data for 555 bp of the mitochondrial gene cytochrome b in plethodontid salamanders, taken from 18 ingroup (tribe Bolitoglossini) and 4 outgroup (tribe Plethodontini) taxa. There were 257 variable sites, of which 219 were phylogenetically informative. Sequence differences among taxa exceeded 20%, and there were up to 15% amino acid differences among the sequences. We also analyzed 37 morphological (including karyological) characters, taken from the literature. Data were analyzed separately and then combined using parsimony and likelihood approaches. There is little conflict between the morphological and DNA data, and that which occurs is at nodes that are weakly supported by one or both of the data sets. Treated separately, the morphological and DNA data provide strong support for some nodes but not for others. The combined data act synergistically so that good support is obtained for nearly all of the nodes in the tree. Recent divergences are supported by silent transitions, and older divergences are supported by a combination of morphological, karyological, DNA transversion, and amino acid changes. Eliminating silent changes from the DNA data improves the consistency index and improves some bootstrap and decay index values for several deeper branches in the tree. However, the combined data set with all characters included provides a better supported tree overall. Maximum likelihood and parsimony with all of the data give not only the same topology but also remarkably similar branch lengths. Results of this analysis support the monopoly of the supergenera Hydromantes and Batrachoseps, and of a sister group relationship of Batrachoseps and the supergenus Bolitoglossa (represented in this study one species of the genus Bolitoglossa).

Animals↗
PubMed1993

Conserved sequence and functional domains in satellite 2 from three families of salamanders.

Thirteen satellite 2 elements from Ambystoma talpoideum and 16 from Amphiuma tridactylum were cloned, sequenced, and compared with the satellite 2 consensus from Notophthalmus viridescens. These elements have maintained a high degree of similarity during the 65-200 Myr that the salamander families, represented by the three species, have been separated. The DNA sequences of the consensus elements from A. talpoideum and A. tridactylum are 81% similar, and both are approximately 65% similar to the N. viridescens consensus. In addition to its DNA sequence, the functional properties of satellite 2 have also been conserved. By selecting and analyzing clones that closely mimicked the consensus of each species, we were able to demonstrate that satellite 2 from each species was capable of promoting transcription after injection into Xenopus laevis oocytes and that synthetic transcripts of satellite 2 from each species were capable of catalyzing their own site-specific cleavage. These properties may be related to the process of retroposition, which was previously proposed to be responsible for the genomic proliferation of satellite 2. Each of these functional properties also has general biological interest.

Ambystoma↗
PubMed1989

Patterns of ribosomal RNA evolution in salamanders.

Sequence comparisons are presented for four segments of the large subunit of ribosomal RNA, including divergent domains D7a and D7b, portions of the large divergent domains D2, D3, and D8, and evolutionarily conservative sequences flanking divergent domains. These results resolve phylogenetic relationships among exemplars of seven families of salamanders and the three amphibian orders. Phylogenetic analysis confirms the prediction that divergent domains feature the highest relative rates of base substitution and length variation within the ribosome, but the divergent domains evolve more slowly than nuclear noncoding DNA and the silent sites of structural genes. Base substitutions demonstrate approximately twice as many transitions as transversions and an uneven distribution among sites within the divergent domains but no apparent bias in base composition. Length mutations are primarily small insertions and deletions, with deletions predominating. The divergent domains appear to be a good source of phylogenetic information for evolutionary events occurring approximately 100-200 million years ago.

Animals↗
PubMed1991

Molecular phylogeny of plethodonine salamanders and hylid frogs: statistical analysis of protein comparisons.

The bootstrapping method of determining confidence in the topology of phylogenetic trees has been applied to electrophoretic protein data for two groups of amphibians: salamanders of two North American genera (Aneides and Plethodon) of the tribe Plethodontini and Holarctic hylid frogs. Some current methods of phylogenetic reconstruction for electrophoretic protein data have been evaluated by comparing the trees obtained from molecular data sets with available morphological data. Molecular data on the phylogenetic relationships of Aneides and Plethodon, data obtained from electrophoretic and immunological studies, indicate that Aneides probably was derived from western Plethodon subsequent to the separation of eastern and western Plethodon. Thus Plethodon very likely is a paraphyletic genus. The extremely low rate of morphological evolution in Plethodon compared with that in Aneides causes difficulty in indicating their evolutionary relationships taxonomically because there are no synapomorphic morphological characters that define either eastern or western Plethodon, whereas there are several for the genus Aneides. Thus molecular data alone probably indicate the evolutionary relationships of the species in these genera. Highton and Larson's (1979) arrangement of species of Plethodon into eight species groups is supported. The topologies of the unweighted pair-group method using arithmetic means (UPGMA) and distance Wagner trees were compared with independent morphological and molecular data on the relationships of the 28 plethodonine species. It was found that UPGMA trees indicate relationships that are more in agreement with other information than are those provided by distance Wagner trees. The use of the bootstrap technique indicates that the topologies of UPGMA trees are better supported statistically than are the topologies of distance Wagner trees. Moreover, different addition criteria produce a variety of distance Wagner trees with different topologies, each with several groupings that are not supported statistically. It is concluded that considerable caution should be used in interpreting the topology of distance Wagner trees. Very similar results were obtained with a second data set on 30 taxa of Holarctic hylid frogs. Trees obtained by the neighbor-joining method are more in agreement with UPGMA phenograms and other data, so this method of phylogenetic reconstruction may be useful to systematists not willing to assume constant rates of evolution.(ABSTRACT TRUNCATED AT 400 WORDS)

Amino Acid Sequence↗
PubMed2001

Calcium channel immunoreactivity in the salamander retina.

This study reports the distribution of the alpha1D and alpha1E calcium channel subunits in the neotenous tiger salamander retina based on immunohistochemical techniques. Confocal and light microscopy were used to localize staining with fluorescently tagged antibodies to alpha1D and alpha1E in cross-sectional and flatmount preparations of retina. Alpha1D-immunoreactivity (alpha1D-IR) was localized to the inner and outer plexiform layers (IPL and OPL, respectively), ganglion cell layer (GCL), and optic fiber layer. Alpha1E-IR was found predominantly in the IPL, with scattered, weak representation in the OPL. Alpha1E-IR was not detected in the GCL or fiber layer. These findings suggest that different alpha1 calcium channel proteins have distinctive distributions in retina, which may reflect their unique and different roles in retinal processing and homeostasis.

Ambystoma↗
PubMed2005

Mechanical vibrations from tadpoles' flapping tails transform salamander's carnivorous morphology.

Some prey or predator organisms exhibit striking rapid morphological plastic changes with distinct morphology under the condition of predator or prey presence. Remote chemicals propagating from the inducing agents are the prevalent induction cues for most examples of induction of distinct morphs. Sonic and visual cues, as well as chemical cues, are known as triggers for induction of behavioural plasticity. Here we show that hydraulic vibration originating from flapping tails of anuran tadpoles is a key cue in relation to induction of a distinct carnivorous morphology, a broad-headed morph, in larval salamander Hynobius retardatus, which is able to efficiently capture and handle prey. This result was further supported by the fact that simple mechanical vibrations of tail-like vinyl fins were able to induce the morph without any biological cues. Induction of the morph triggered by hydraulic vibration provides a novel concept for understanding the proximate mechanisms of induction of morphological changes.

Adaptation, Physiological↗
PubMed1981

Merkel cells as targets of the mechanosensory nerves in salamander skin.

Previous investigations showed that the morphological basis of the low-threshold rapidly adapting mechanoreceptors in the salamander skin is neurite-Merkel cell complex located in the epidermis. We have now examined whether sensory nerves are required for the appearance of Merkel cells, and whether these cells act as specific targets for ingrowing sensory axons. Electronmicroscopic examination of denervated skin shows that Merkel cells survive in normal density and appearance in the epidermis for at least six months without nerves. In addition, mechanosensory function of such skin is restored when the skin become reinnervated either by collaterally sprouting (intact) nerves or by regenerating ones, and Merkel cells are then found to have nerve terminals associated with them. These experiments suggest that Merkel cells do act as targets for ingrowing nerves in these circumstances. Further experiments were done on skin that had regenerated in the total absence of any innervation to the limb; such new skin contained nerve-free Merkel cells that were normal in appearance, density and location. When nerves were allowed to innervate this new skin the sensory axons grew to the "new" Merkel cells and established typical neurite-Merkel cell complexes, and normal mechanosensory function. Since the new skin had developed without nerves, the sensory axons were not following "old" pathways to the Merkel cells. We conclude that the Merkel cells are the true targets for the mechanosensory axons; these cells attract the sensory axons to them and/or contain specific markers that allow the ingrowing axons to recognize them. Once a Merkel cell becomes innervated it loses this "target cell" quality. In addition, mechanosensory function of the sensory endings appears to occur only when the endings have developed a morphological association with a Merkel cell.

Animals↗
PubMed1981

Competitive reinnervation of salamander skin by regenerating and intact mechanosensory nerves.

We have investigated in the salamander the possibility that regenerating mechanosensory nerves might prefer the epidermal Merkel cells (their specific targets) that are located within their segmental domain to those within a "foreign" domain. Since regerating nerves cross domain boundaries with no evidence of the marked delay exhibited by intact sprouting nerves, we examined situations in which the regenerating axons of one segmental nerve were effectively in equal competition for denervated skin with those of another segmental nerve. Additionally, we investigated whether there were differences between regenerating axons and intact sprouting axons of the same segmental nerve, in their ability to innervate available skin both inside and outside the parent domain. No preference was detected of any type of nerve, regenerating or intact, for particular skin regions, or for Merkel cells as indicated by the numbers of mechanosensory thresholds of the touch spots that developed in reinnervated skin. Neither was there any indicating of displacement of "foreign" nerves from a particular region by appropriate axons. When regenerating and intact (sprouting) axons invaded denervated skin more or less simultaneously, the former appeared to have a slight advantage since a significantly greater proportion of skin was innervated by regenerated fibres. With this one exception, all the results were explained most simply by assuming that the axon that first arrives at a denervated Merkel cell establishes a permanent association with that cell and at the same time causes it to lose its "target character" for other axons.

Ambystoma↗
PubMed1986

Photoreceptors of the larval tiger salamander retina.

Six morphological types of photoreceptor were identified with electron microscopy in radial sections of the retina of the larval tiger salamander, Ambystoma tigrinum. In order of predominance these six types are: red rods, large single cones, double cones composed of principal and accessory members, small single cones, and green rods. The different types of photoreceptor can be distinguished by a number of morphological and cytological characteristics. The identification of the small single cone type now provides evidence for more than one type of single cone in an amphibian retina. The interconnections of the different types of photoreceptor by gap junctions were studied in tangential sections. Rod-rod and rod-cone gap junctions occurred in all possible combinations, but no cone-cone junctions were found even between members of double cones.

Ambystoma↗
PubMed1992

Voltage-dependent block by L-cis-diltiazem of the cyclic GMP-activated conductance of salamander rods.

Block by L-cis-diltiazem of the cyclic GMP-activated conductance was studied in excised inside-out patches from the salamander rod outer segment. When L-cis-diltiazem was applied from the cytoplasmic face of the patch, current suppression increased monotonically with membrane depolarization, the ratio of blocked to unblocked current varying e-fold in 50 mV. This suggests that L-cis-diltiazem interacts with a binding site located about half-way across the membrane field, and is unable to fully traverse the cyclic GMP-activated channel. The kinetics of block were accelerated by increasing L-cis-diltiazem concentration and by depolarization. These results can be fitted by a single barrier model in which the barrier peak is located about a third of the way across the membrane field from the cytoplasmic face. Application of L-cis-diltiazem from the extracellular face of the patch also resulted in an enhancement of block with membrane depolarization. Indirect evidence supports the notion that this block resulted from partition of the unchanged form of the blocker across the membrane, and its subsequent interaction with the cytoplasmic face of the conductance.

Ambystoma↗
PubMed1994

The effect of pH on the block by L-cis-diltiazem and amiloride of the cyclic GMP-activated conductance of salamander rods.

Block by L-cis-diltiazem and amiloride of the cyclic GMP-activated conductance was studied in inside-out patches excised from the salamander rod outer segment. When cytoplasmic pH was varied, the steady-state level of block by L-cis-diltiazem changed in the way that would be predicted if it were the protonated ammonium group that is responsible for effecting block. This is in contrast to the recent results of Haynes (J. gen. Physiol. 100, 783 (1992)) in catfish cones, where no such change was seen. Amiloride was found to block the conductance with a similar voltage dependence to that of L-cis-diltiazem. The dependence of amiloride block on cytoplasmic pH was found to be shifted relative to that of L-cis-diltiazem, consistent with the 1 pH-unit higher pKa value of amiloride and the idea that it is only the charged form of amiloride which can effect block. This suggests that the results seen with L-cis-diltiazem were indeed due to changes in the proportion of blocker in the protonated form, and not to effects of protons on the channel.

Ambystoma↗