Epidermal keratinization in the salamander and a comparison with other amphibia.
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Relative size and arrangement of the brain and paired sense organs are examined in three species of Thorius, a genus of minute, terrestrial salamanders that are among the smallest extant tailed tetrapods. Analogous measurements of representative species of three related genera of larger tropical (Pseudoeurycea, Chiropterotriton) and temperate (Plethodon) salamanders are used to identify changes in gross morphology of the brain and sense organs that have accompanied the evolution of decreased head size in Thorius and their relation to associated changes in skull morphology. In adult Thorius, relative size (area measured in frontal plane, and length) of the eyes, otic capsules, and brain each is greater than in adults of all of the larger genera; relative size of the nasal capsules is unchanged or slightly smaller. Interspecific scaling phenomena--negative allometry of otic capsule, eye and brain size, isometry or slight positive allometry of nasal capsule size, all with respect to skull length--also are characteristic of intraspecific (ontogenetic) comparisons in both T. narisovalis and Pseudoeurycea goebeli. Predominance of the brain and eyes in Thorius results in greater contact and overlap among these structures and the nasal capsules in the anterior portion of the head. This is associated with anterior displacement of both the eyes and nasal capsules, which now protrude anterior to the skull proper; a change in eye shape; and medial deformation of anterior braincase walls. Posteriorly, predominance of the otic capsules has effected a reorientation of the jaw suspensorium to a fully vertical position that is correlated with the novel presence of a posteriorly directed squamosal process and shift in origin of the quadropectoralis muscle. Many of these changes in cranial morphology may be explained simply as results of mechanical (physical) interactions among the skeletal, nervous, and sensory components during head development at reduced size. This provides further evidence of the role of nervous, sensory, and other "soft" tissues in cranial skeletal morphogenesis, and reinforces the need to consider these tissues in analyses of skull evolution.
The morphology and fine structure of the basilar recess and basilar papilla were investigated in four species of salamanders from the family Ambystomatidae. The otic relationships of the recess and papilla to the proximal part of the lagena and saccule are described, and new terminology is suggested for the periotic relationships of the basilar recess to a diverticulum of an intracapsular periotic sac. The basilar papilla consists of supporting cells united laterally by gap junctions, capped by microvilli uniformly arranged around a short, central cilium, and hair cells that typically show several synapses with a single afferent nerve fiber, each marked by a rounded synaptic body surrounded by vesicles. In contrast to anuran basilar papillae, efferent nerve terminals were observed in synapse with hair cells and, rarely, upon afferent fibers. The distal half of the ambystomatid papilla contained hair cells capped by tall ciliary bundles, with kinocilia that show swellings near their tips with delicate attachments to adjacent tall stereocilia. A tectorial body covers only this region of the papilla. Hair cells with shorter stereocilia, situated in the proximal half and at the papillar margins, are related only to filamentous extensions of the tectorial body. The ambystomatid basilar recess and papilla are compared to auditory end-organs in other vertebrates, and it is suggested that a basic distinction can be made between aural neuroepithelia in amniotes versus that in nonamniotic vertebrate ears.
The surface morphology of the basilar recess and papilla was examined in 14 species of newts and salamanders selected from the five families of urodeles (Ambystomatidae, Salamandridae, Hynobiidae, Cryptobranchidae, and Amphiumidae) known to have this end-organ. In this sampling, the general organization of basilar structures is essentially similar across species investigated. The recess forms a tubular diverticulum of the proximal part of the lagena. One wall of the recess is associated with a diverticulum of the intracapsular periotic sac, and an adjacent wall is occupied by the basilar papilla. The papilla contained from as few as five hair cells in specimens of Taricha torosa to over 200 hair cells in Cryptobranchus allegheniensis. In most species, the papilla showed a morphological continuum between tall centrally or distally placed ciliary bundles and short ciliary bundles near the papillar margins. In certain species examined, tall bundles had kinocilia with swellings near their tips. Most forms showed a tendency to have groups of ciliary bundles morphologically polarized either toward or away from the saccule. In Cryptobranchus and Dicamptodon, many bundles had a random orientation. The gross and fine structural features of the basilar complex are compared in urodeles and anurans, and "generalized" features for the amphibian basilar complex are suggested. The basilar complex of Cryptobranchus is interpreted as being most generalized, representing a structural form from which most features of the basilar complex in other urodeles and anurans can be derived.
The innervation of the musculature of the tongue and the hyobranchial apparatus of caecilians has long been assumed to be simple and to exhibit little interspecific variation. A study of 14 genera representing all six families of caecilians demonstrates that general patterns of innervation by the trigeminal, facial, glossopharyngeal, and vagus nerves are similar across taxa but that the composition of the "hypoglossal" nerve is highly variable. Probably in all caecilians, spinal nerves 1 and 2 contribute to the hypoglossal. In addition, in certain taxa, an "occipital," the vagus, and/or spinal 3 appear to contribute fibers to the composition of the hypoglossal nerve. These patterns, the lengths of fusion of the contributing elements, and the branching patterns of the hypoglossal are assessed according to the currently accepted hypothesis of phylogenetic relationships of caecilians, and of amphibians. An hypothesis is proposed that limblessness and a simple tongue, with concomitant reduced complexity of innervation of muscles associated with limbs and the tongue, has released a constraint on pattern of innervation. As a consequence, a greater diversity and, in several taxa, greater complexity of neuroanatomical associations of nerve roots to form the hypoglossal are expressed.
Histology of the cloacae of Rhyacotriton olympicus and representative species from the genera Ambystoma and Dicamptodon was examined by light microscopy. Females of Ambystoma possess sperm storage glands, the spermathecae, as well as ventral glands and dorsal glands, both of uncertain function. Females of Ambystoma examined from the subgenus Linguaelapsus differ from those in the subgenus Ambystoma by possessing more extensive ventral gland clusters and a shorter cloacal tube. Females of Dicamptodon possess spermathecae and ventral glands, but differ in cloacal conformation from females of Ambystoma and lack the dorsal glands. Females of R. olympicus possess more extensive epidermal lining in the cloaca than that found in females of Ambystoma and Dicamptodon, and the only glands present are spermathecae, which cluster around a tube in the dorsal roof. Males of Ambystoma, Dicamptodon, and R. olympicus possess five types of cloacal glands (dorsal pelvic glands, lateral pelvic glands, anterior ventral glands, posterior ventral glands, and Kingsbury's glands) that function in spermatophore formation, and vent glands that may produce a courtship pheromone. In Ambystoma and Dicamptodon, vent glands secrete along the medial borders of the cloacal orifice. Males of A. opacum and A. talpoideum differ from males of other species examined from the two genera by possessing more extensive vent glands. Males of R. olympicus possess unique vent glands in which tubules secrete onto the surface of vent lobes lateral to the posterior end of the cloacal orifice, and distal ends of the glands pass anteriorly, superficial to the fascia enclosing the other cloacal glands. The results from analysis of cloacal anatomy support other data indicating that Ambystoma and Dicamptodon are sister groups, and that Rhyacotriton olympicus is not closely related to either of the other two genera and merits placement in a separate family.
Microcorrosion casts of the renal vascular system of tadpoles of the Clawed Frog, Xenopus laevis, were observed by scanning electron microscopy. Glomerular differentiation was studied qualitatively and quantitatively during developmental stages 56-66 (metamorphic climax). The general structure of the renal vascular system corresponds to the pattern commonly found in anurans; however, the arterial supply has conspicuous connecting vessels that supply groups of glomeruli. In the dorsal part of the kidney, qualitative differentiation of glomerular structures precedes quantitative growth. The ventral part of the kidney has larger, well-developed renal corpuscles of nearly adult appearance. Four developmental stages of glomerulogenesis are distinguished morphologically and their glomerular and vascular growth is analyzed.
Sperm storage glands, spermathecae, were examined from mated female Ambystoma opacum during the breeding season. No differences occur in the spermathecal ultrastructure of individuals sacrificed prior to oviposition and those sacrificed within 3 days of removal from tended clutches of recently oviposited eggs. The simple tubuloalveolar glands produce two types of secretory vacuoles. Apical secretory vacuoles contain glycosaminoglycans for export into the lumen to bathe stored sperm, perhaps providing the chemical/osmotic environment necessary for sperm quiescence. The other type of secretory vacuole contains an unsaturated lipid that is produced for export into the connective tissue surrounding the spermathecae. The role of this secretion may involve the contraction of myoepithelial cells, resulting in sperm expulsion. Some sperm undergo degradation in the spermathecal epithelium, and an interepithelial leukocyte was observed in one specimen. Apical secretory vacuoles and sperm are absent from the spermathecae of a specimen sacrificed 62 days after removal from a tended egg clutch. This is the first report on the spermathecal cytology of a salamander from the Ambystomatidase, and comparisons with salamanders from other families provide a morphological basis for considering spermathecae polyphyletic within the Caudata.
The skin of the aquatic pipid frog, Xenopus laevis, was examined for specific biomechanical features: 1) thickness, 2) maximal strain at break (epsilon f), 3) tensile strength (sigma m), 4) modulus of elasticity (E, stiffness), and 5) the area under the stress-strain curve (W) (breaking energy, toughness). Skin freshly removed from dorsal, ventral, and lateral areas of the body was subjected to uniaxial tension. In both sexes, the dorsal skin is thicker than the ventral. The skin of male frogs was consistently thinner in all body regions than that of females. Most biomechanical parameters showed a considerable range of values in both males (epsilon f = 59-63%, sigma m = 15-16.5 MPa, E = 33.5-38.4 MPa, W = 3.8-4.5 MJ/m3) and females (epsilon f = 102-126%, sigma m = 11.5 MPa, E = 10.4-12 MPa, W = 5.2-6.7 MJ/m3). The disparate epsilon f values in males (low) and females (high) might reflect sexual dimorphism. Static stress-strain curves were typically J-shaped; with the exception of a "toe," the curves rose approximately linearly with increasing strain. The skin of X.laevis, although heterogeneous in structure, possesses features similar to those found in tissues with aligned collagen fibers such as tendons or fish skin. However, in anurans, the skin seems to play a more passive mechanical role during locomotion than in fish.
The larval neurocranium and visceral arches of seven dendrobatid species representing four genera are described, based on cleared-and-stained and serially sectioned specimens. A variety of characters is shared by all seven species. Larval features do not substantiate the assumption of close ranoid affinities of the Dendrobatidae. Instead dendrobatid larvae share features such as the special quadripartite cartilago suprarostralis, the lack of the larval processus oticus, the presence of three foramina acustica, and the lack of a foramen perilymphaticum accessorius with many bufonoid larvae. The first of these characters is unique to bufonids, hylids, dendrobatids, and some New World leptodactylids; the other characters also occur in pelobatids and are presumably plesiomorphic for the Neobatrachia. The free proximal ends of Ceratobranchialia II and III are an autapomorphy of the Dendrobatidae supporting the monophyly of the family. Some features of the cranium are paedomorphic: low cartilago orbitalis, lack of connection between cartilage orbitalis and otic capsule (most species), and vestigal taeniae tecti. New anatomical terms are introduced.
The advertisement call of the leptodactylid frog Batrachyla antartandica from southern Chile consists of a train of brief percussive tone pulses whose energy is centred at about 2 kHz. To gain an understanding of the temporal features that are essential for call recognition, playback experiments were conducted with 11 males. Subjects were presented with a synthetic imitation of this signal and variants for which different temporal call parameters were modified systematically. The number of pulses, pulse rate and latency of evoked vocal responses (EVRs) to stimuli having high pulse repetition rates (i.e. 8 and 16 pulses/s) were significantly weaker relative to responses to stimuli having an equal number of pulses but lower pulse rates. A similar, non-significant tendency was observed for a series of stimuli with different pulse rates for which the total stimulus duration was held constant. EVRs also decreased significantly for stimuli having long pulse durations (i.e. 48 and 96 ms) relative to stimuli comprising shorter pulses. No significant differences were observed between EVRs to stimuli for which pulse rise and fall times were varied from 1-20 ms. Responses to calls comprising trains of 10 pulses were weaker compared with stimuli having fewer pulses per train. The selective EVRs of B. antartandica for different temporal parameters contributes to an understanding of the mechanisms involved in call recognition and stress the relevance of temporal processing of sound by males for the emergence of specific patterns of vocal behaviour in anurans.Copyright 1997 The Association for the Study of Animal Behaviour1997The Association for the Study of Animal Behaviour
Estrogens are involved in gonadal differentiation in birds and in reptiles with temperature-dependent sex determination, suggesting a key role for aromatase, the enzyme complex converting androgens to estrogens. Pleurodeles waltl is an amphibian with ZZ/ZW genotypic sex determination but gonadal differentiation is thermosensitive. The sexual phenotype is in conformity with the sexual genotype at ambient temperature (20 +/- 2 degrees), but ZW animals are sex-reversed in functional males when larvae are reared at 32 degrees from stage 42 to stage 54 (thermosensitive period). Histological sexual differentiation of gonads begins at stage 53. Aromatase activity was measured either in the gonadmesonephric complexes (stages 47 to 52) or in the gonads (stages 53 to 56) of ZZ and ZW larvae reared at ambient temperature or at 32 degrees and of ZW larvae shifted at stages 53, 55, or 56 from ambient temperature to 32 degrees for 48 hr. At ambient temperature, aromatase activity was detected, at low levels, in gonad-mesonephric complexes of both ZZ and ZW larvae at stages 47 and 50. At stage 52, it was significantly higher in ZW individuals than in ZZ ones. Then it remained low in gonads of ZZ males, whereas it markedly increased in gonads of ZW females. At 32 degrees, the gonad-mesonephric complexes (stage 52) and the gonads (stages 53 to 56) of ZW larvae had low aromatase activity similar to that in ZZ individuals. The exposure to 32 degrees for 48 hr of ZW individuals significantly decreased gonadal aromatase activity during the thermosensitive period (shift at stage 53) but not after the thermosensitive period (shifts at stages 55 and 56).(ABSTRACT TRUNCATED AT 250 WORDS)
Insulin has been isolated from an extract of the pancreas of a salamander, the lesser siren Siren intermedia, and its primary structure was established as: A-chain, Gly-Ile-Val- Glu-Gln-Cys-Cys-His-Asn-Thr10-Cys-Ser-Leu-Tyr-Gln-Leu-Glu-Asn-Tyr- Cys20-Asn, and B-chain, Val-Pro-Asn-Lys-Pro- Leu-Cys-Gly-Ala-His10-Leu-Val-Glu-Val-Met-Tyr-Phe-Val- Cys-Gly20-Asp-Arg-Gly-Phe-Phe-Tyr-Pro-Ser-Ser-Thr 30. Although those amino acid residues considered to constitute the receptor-binding region of insulin have been retained, siren insulin contains several substitutions (Gln-->Lys at B4, Ser-->Ala at B9, Ala-->Val at B14, Leu-->Met at B15, Leu-->Phe at B17, Pro-->Ser at B28, and Lys-->Ser at B29) of amino acid residues that are conserved in insulins from species of other amphibian orders. The biological activity of siren insulin was not investigated in this study but the substitutions at B28 (involved in dimer formation) and at B14 and B17 (involved in hexamer formation) may be expected to influence conformation and therefore biological potency. The data are consistent with the view that the Sirenoidea represent an early divergence from the ancestral stock of salamanders.
We investigated the relationships of Asian bufonids using partial sequences of mitochondrial DNA genes. Twenty-six samples representing 14 species of Bufo from China and Vietnam and 2 species of Torrentophryne from China were examined. Three samples of Bufo viridis from Armenia and Georgia were also sequenced to make a comparison to its sibling tetraploid species B. danatensis. Bufo americanus, from Canada, was used as the outgroup. Sequences from the 12S ribosomal RNA, 16S ribosomal RNA, cytochrome b, and the control region were analyzed using parsimony. East Asian bufonids were grouped into two major clades. One clade included B. andrewsi, B. bankorensis, B. gargarizans, B. tibetanus, B. tuberculatus, its sister clade B. cryptotympanicus, and the 2 species of Torrentophryne. The second clade consisted of B. galeatus, B. himalayanus, B. melanostictus, and a new species from Vietnam. The placement of three taxa (B. raddei, B. viridis, and its sister species, B. danatensis) was problematic. The genus Torrentophryne should be synonymized with Bufo to remove paraphyly. Because B. raddei does not belong to the clade that includes B. viridis and B. danatensis, it was removed from the viridis species group. The species status of B. bankorensis from Taiwan is evaluated.
An analysis of partial sequences of the 16S ribosomal rRNA gene (582 bp) of 20 poison frog species (Dendrobatidae) confirmed their phylogenetic relationships to bufonid and leptodactylid frogs. Representatives of the ranoid families and subfamilies Raninae, Mantellinae, Petropedetinae, Cacosterninae, Arthroleptidae, Astylosternidae, and Microhylidae did not cluster as sister group of the Dendrobatidae. Similar results were obtained in an analysis using a partial sequence of the 12S gene (350 bp) in a reduced set of taxa and in a combined analysis. Within the Dendrobatidae, our data supported monophyly of the genus Phyllobates but indicated paraphyly of Epipedobates and Colostethus. Minyobates clustered within Dendrobates, contradicting its previously assumed phylogenetic position. Phobobates species clustered as a monophyletic unit within Epipedobates. Allobates was positioned in a group containing two Colostethus species, indicating that lack of amplexus, presence of skin alkaloids, and aposematic coloration evolved independently in Allobates and the remaining aposematic dendrobatids.
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Specialized cells are generally unable to switch from one type to another. For example, muscle cells can not be converted to intestine, brain, skin, etc. However, the nucleus of a specialized cell, such as muscle, can be transplanted to an enucleated egg, and the resulting combination will form all these and most other cell types. Likewise somatic cell nuclei injected into oocytes will switch their pattern of gene expression to conform to that of an oocyte, repressing some genes and activating others. This article reviews what is known of the mechanisms involved in these examples of nuclear reprogramming.