两栖动物性染色体的多样性及其进化机制的研究进展

两栖动物性别决定类型和性染色体具有多样性的特点。在已发现异形性染色体两栖动物中,大部分物种Y或W染色体大于其对应的X或Z染色体,少数物种具有高度分化的Y或W染色体。同时两栖动物类群内基因组大小差异大,性染色体间分子水平上也存在差异。高频转换、偶然重组和染色体重排可能是两栖动物性染色体进化过程中的关键机制。本综述通过对两栖动物性染色体进化的深入探讨,揭示其遗传性别决定的机理,有助于对两栖动物性别人工调控的进一步探索。     

The evolution of metamorphosis in amphibians

A survey is provided of the external transformations that coincide with metamorphosis or a water-to-land transition, and of transformations during water-to-land transition in the retinal projection, the brain stem, the lateral-line system, and the inner ear of amphibians. Among the three orders of amphibians, the frogs are characterized by more pronounced transformations during the water-to-land transition than are the other two orders. Some of the progressive and regressive changes in the sensory and nervous system are presented and a scenario is suggested for the evolution of these transformations among amphibians. Suggestions that metamorphosis in frogs can recapitulate the water-to-land transition of ancestral amniotic vertebrates are refuted.     

Phylogenetic Comparison of the Photoaffinity-Labeled Benzodiazepine Receptor Subunits

The late evolutionary appearance of the benzodiazepine receptor (BZR) allows an experimental approach for evaluation of the qualitative development of its subunits. Photoaffinity labeling of brain membranes with [3H]flunitrazepam followed by sodium dodecyl sulfate-polyacrylamide gel electrophoresis and fluorography offers a suitable method for tracing the qualitative evolution of the BZR. A systematic comparison of the subunit patterns in fishes, amphibians, reptiles, birds, and mammals revealed that the subunit of 53K is phylogenetically the oldest photoaffinity labeled subunit; whereas it is the only band present in the lungfish and most amphibians, additional bands are apparent in higher tetrapods. In fishes, the evolution of the BZR subunits leads to the loss of the 53K subunit. KD values are discussed in relation to specific subunit patterns. Possible explanations for the observed variation of the subunits are discussed, with special emphasis placed on the possible evolution by gene duplication and subsequent divergence.     

Circadian Rhythms in Amphibians and Reptiles: Ecological Implications

Circadian rhythms of amphibians and reptiles in the field and under semi-natural conditions and the underlying mechanisms, including the ways of entrainment to environmental cues and the oscillators driving the rhythms, have been reviewed. Studies on the behavioral rhythms in the field are meager in both amphibians and reptiles. In anuran amphibians, Xenopus adults showed more robust nocturnal locomotor activity than did tadpoles. This indicates the ecological significance of the differences in activity pattern shown by amphibians at different life stages, because differences between adults and young in the same environment may serve to isolate partially the young from the adults' cannibalism. In reptiles, free-running rhythms are more robust and continue for a longer time compared to amphibians. In both amphibians and reptiles, multi-photoreceptors are involved in photo-entrainment of circadian rhythms. The eyes, pineal complex and deep brain comprise a multi-oscillator system as well as a multi-photoreceptor system.     

Metallothionein primary structure in amphibians: Insights from comparative evolutionary analysis in vertebrates

Metallothioneins are cysteine-rich, low-molecular weight metal-binding proteins ubiquitously expressed in living organisms. In the last past years, the increasing amount of vertebrate non-mammalian metallothionein sequences available have disclosed for these proteins differences in the primary structure that have not been supposed before. To provide a more up-to-date view of the metallothioneins in non-mammalian tetrapods, we decided to increase the still scarce knowledge concerning the primary structure and the evolution of metallothioneins in amphibians. Our data demonstrate an unexpected diversity of metallothionein sequences among amphibians, accompanied by remarkable features in their phylogeny. Phylogenetic analysis also reveals the complexity of vertebrate metallothionein evolution, made by both ancient and more recent events of gene duplication and loss.     

Neurobiological mechanisms of the origin of terrestrial vertebrates

Some trends in the phylogenetic changes of the vertebrate neural system in the course of their transition from aquatic to terrestrial mode of life are analyzed. Wide multifunctionality of the neural centers and heterotopic substitution of the brain functions are supposed to be prerequisites of transition of archaic amphibians to the land via a stable transitional environment. The latter include underground or vegetation aquatic-air labyrinths in which conditions of gradual acquisition of adaptations to the terrestrial mode of life occurred. The principal neurological adaptations to such mode of life and less dependence on aquatic environment developed during the long-term evolution in that environment. The neurobiological model of evolutionn of the archaic amphibians in the transitional environment of aquatic-air labyrinths suggested herewith is compared to other hypotheses of the tetrapod origin.     

Left and right in the amphibian world: which way to develop and where to turn?

The last decade has seen a dramatic increase in studies on the development, function and evolution of asymmetries in vertebrates, including amphibians. Here we discuss current knowledge of behavioral and anatomical asymmetries in amphibians. Behavioral laterality in the response of both adult and larval anurans to presumed predators and competitors is strong and may be related, respectively, to laterality in the telencephalon of adults and the Mauthner neurons of tadpoles. These behavior lateralities, however, do not seem to correlate with visceral asymmetries in the same animals. We briefly compare what is known about the evolution and development of asymmetry in the structure and function of amphibians with what is known about asymmetries in other chordate and non-chordate groups. Available data suggest that the majority of asymmetries in amphibians fall into two independent groups: (1) related to situs viscerum and (2) of a neurobehavioral nature. We find little evidence linking these two groups, which implies different developmental regulatory pathways and independent evolutionary histories for visceral and telencephalic lateralizations. Studies of animals other than standard model species are essential to test hypotheses about the evolution of laterality in amphibians and other chordates.     

Recent Progress in Understanding Early Tetrapods

This paper reviews significant discoveries and interpretationsmade for Paleozoic tetrapods over the past twenty-five years.In that span twelve significant, new localities have been found,including the oldest ever at about 370 million years in age.About 60 new genera have been described; five providing importantinsight into the early evolution of land vertebrates. The numberof exceptionally well known taxa with multiple, excellentlypreserved specimens has doubled to eight from four. The veryearliest tetrapods have been discovered to have been polydactylous,the ear region to have had a complicated early evolution andthe specialized tooth type found in Recent amphibians has beenfound in a group of Lower Permian temnospondyl amphibians, indicatingan evolutionary relationship. Perhaps the most significant advancein understanding the evolution of early tetrapods is that thebasal amniote groups have become better characterized and astart has been made in providing a defensible hypothesis fortheir relationships. The ascendancy of cladistics, functionalmorphology and plate tectonics has changed the way paleontologistsview fossils resulting in more defensible phylogenies and behavioraland biogeographical scenarios. These approaches to understandinghave, perhaps, had a more profound impact than any of the newfossil discoveries.     

Large‐brained frogs mature later and live longer

Brain sizes vary substantially across vertebrate taxa, yet, the evolution of brain size appears tightly linked to the evolution of life histories. For example, larger brained species generally live longer than smaller brained species. A larger brain requires more time to grow and develop at a cost of exceeded gestation period and delayed weaning age. The cost of slower development may be compensated by better homeostasis control and increased cognitive abilities, both of which should increase survival probabilities and hence life span. To date, this relationship between life span and brain size seems well established in homoeothermic animals, especially in mammals. Whether this pattern occurs also in other clades of vertebrates remains enigmatic. Here, we undertake the first comparative test of the relationship between life span and brain size in an ectothermic vertebrate group, the anuran amphibians. After controlling for the effects of shared ancestry and body size, we find a positive correlation between brain size, age at sexual maturation, and life span across 40 species of frogs. Moreover, we also find that the ventral brain regions, including the olfactory bulbs, are larger in long‐lived species. Our results indicate that the relationship between life history and brain evolution follows a general pattern across vertebrate clades.     

The amphibian octavo-lateralis system and its regressive and progressive evolution

The phylogenetic and ontogenetic changes in the octavolateralis system of sarcopterygian fish and tetrapods, presumed to be important for the formation of an amphibian auditory system, are reviewed. The lateral line system shows rudimentation of lines and loss of ampullary electroreceptors in many amphibians; in some amphibians it never develops. The metamorphic changes of the lateral-line system show different patterns in the different amphibian lineages with metamorphic retention in most urodeles and metamorphic loss in most anurans. The multitude of both ontogenetic and phylogenetic changes of the lateral line system among amphibians do exclude any prediction as to how this system might have changed in ancestral amniotes. The most important auditory epithelium of the tetrapod inner ear, the basilar papilla, seems to be primitively present in all tetrapods and Latimeria. In two amphibian lineages there is a trend towards rudimentation and loss of the basilar papilla. Only in the third order, the anurans, a tympanic ear develops and the inner ear shows a progressive evolution of the auditory epithelia. Together with the known differences in the periotic labyrinth of amphibians and amniotes, this scenario suggests a parallel evolution of the amniotic and anuran auditory periphery. All mechanoreceptive hair cells of the lateral line system and the inner ear appear to receive a common and bilateral efferent innervation. Among amphibians this pattern is represented only in some urodeles, whereas anurans show a derived pattern with loss of a bilateral component and presumably also of a common neuromast/inner ear component. Changes in the rhombencephalic nuclei which receive octavo-lateralis afferent fibers show a trend towards development of auditory nuclei only in the anuran lineage. The phylogenetic appearance of an auditory nucleus in this lineage coincides with the complete absence of formation of ampullary electroreceptors. In contrast, the earlier claim of a correlation between a metamorphic loss of the lateral line system and the formation of an auditory nucleus is not supported by more recent data: an auditory nucleus develops in anurans already prior to metamorphosis and is present in all anurans even when they retain the neuromast system. In anurans with a metamorphic loss of the neuromasts, the second order neurons degenerate as well. This independence of the auditory and the second order lateral line nuclei is further substantiated by their separate projection to other brain areas, like the torus semicircularis of the midbrain, and their functional properties.(ABSTRACT TRUNCATED AT 400 WORDS)     

Life history of amphibians and gravity.

Anurans hold a unique position in vertebrate phylogeny, as they made the major transition from water to land. Through evolution they have acquired fundamental mechanisms to adapt to terrestrial gravity. Such mechanisms are now shared among other terrestrial vertebrates derived from ancestral amphibians. Space research, using amphibians as a model animal, is significant based on the following aspects: (1) Anuran amphibians show drastic changes in their living niche during their metamorphosis. Environments for tadpoles and for terrestrial life of frogs are quite different in terms of gravity and its associated factors. (2) Certain tadpoles, such as Rhacophorus viridis amamiensis, have a transparent abdominal wall. Thus visceral organs and their motion can be observed in these animals in non-invasive manner through their transparent abdominal skin. This feature enables biologists to evaluate the physiological state of these amphibians and study the autonomic control of visceral organs. It is also feasible for space biologists to examine how such autonomic regulation could be altered by microgravity and exposure to the space environment.     

Cranial kinesis in the amphibia: a review

All extant orders of amphibians are characterized by kinetic skulls. Main type of intracranial movability in amphibians is pleurokinetism, that is supplemented in different amphibian groups by various types of rhyncho- and prokinetism. The most primitive pattern of cranial kinesis is revealed in the stegocrotaphic gymnophions. More paedomorphic species retain general cranial flexibility that is characteristic of larval skull. That is unfavourable for evolution of well-regulated (adult) cranial kinesis and related feeding adaptations. Kinetism is also reduced in the species with heavily ossified skulls. Adaptive role and evolution of cranial kinesis in amphibians are discussed.     

Brain, Gut and Skin Peptide Hormones in Lower Vertebrates

Understanding of peptide hormone evolution rests primarily onstructural information, either direct or inferred. We summarizestudies of fishes and amphibians to provide initial informationwithin the vertebrate lineage for selected peptides which exhibitvarying structural heterogeneity. For these peptides, thyrotropin-releasinghormone, somatostatin, luteinizing hormone-releasing hormoneand cholecystokinin related peptides manifest increasing diversification.Members of these peptide families are found distributed amonga variety of tissues (e.g., brain, gut, skin, retina, sympatheticnervous system), yet the number of genes encoding for individualtypes of peptides is presently uncertain. We emphasize the needfor additional structural information, for a more thorough anddiverse taxonomic investigation within the vertebrate lineage,and for specification of those genetic elements which ultimatelydetermine evolutionary opportunities for peptide evolution.     

Evolution of anuran brains: disentangling ecological and phylogenetic sources of variation

Variation in ecological selection pressures has been implicated to explain variation in brain size and architecture in fishes, birds and mammals, but little is known in this respect about amphibians. Likewise, the relative importance of constraint vs. mosaic hypotheses of brain evolution in explaining variation in brain size and architecture remains contentious. Using phylogenetic comparative methods, we studied interspecific variation in brain size and size of different brain parts among 43 Chinese anuran frogs and explored how much of this variation was explainable by variation in ecological factors (viz. habitat type, diet and predation risk). We also evaluated which of the two above‐mentioned hypotheses best explains the observed patterns. Although variation in brain size explained on average 80.5% of the variation in size of different brain parts (supporting the constraint hypothesis), none of the three ecological factors were found to explain variation in overall brain size. However, habitat and diet type explained a significant amount of variation in telencephalon size, as well in three composite measures of brain architecture. Likewise, predation risk explained a significant amount of variation in bulbus olfactorius and optic tecta size. Our results show that evolution of anuran brain accommodates features compatible with both constraint (viz. strong allometry among brain parts) and mosaic (viz. independent size changes in response to ecological factors in certain brain parts) models of brain size evolution.     

A genetic model for the evolution of the glycosphingolipids

The glycosphingolipids have been found in many animal tissues, but the complexity of their molecular structure varies considerably among the different phyla. Relatively simple structures have been found in invertebrate species, while the most complex have been demonstrated in brain tissue of modern fishes and amphibians. The data on the phylogenetic distribution of the glycosphingolipids has been interpreted to indicate that a significant number of gene duplications, involving many different structural genes, may have occurred during a few specific periods of vertebrate evolution. The transition from invertebrate to jawless vertebrate, the divergence of rays and skates from true sharks, the advent of modern bony fishes and the transition from aquatic to terrestrial vertebrates, each could have veen accompained by duplications of genes involved in the synthesis and degradation of glycosphingolipids. The evolutionary study of such a multi-enzyme system may be one means to detect alterations in the genome as a whole. The apparent correspondence in time of these gene duplications involved in glycosphingolipid metabolism and periods of rapid vertebrate evolution which may have been accompanied by significant increases in the amount of cellular DNA suggests that such changes may have occurred via the mechanism of tetraploidization.     

The C. U. Ariëns Kappers brain collection

The C. U. Ari?ns Kappers brain collection, at the Netherlands Institute for Neuroscience in Amsterdam, is one of the largest and oldest of the world's catalogued repositories of specimens that reveal the course of brain evolution and the resulting panoply of neural biodiversity. Established a century ago, it has served since then as the basis of the encyclopedic texts authored by its founder, as well as research publications into the current time. It consists of 726 specimens: these include 309 mammals, 134 birds, 81 reptiles, 21 amphibians, and 179 "pisces"--a grouping of bony fish, sharks, and cyclostomes. We present here accounts of the history and contents of this treasure trove of research materials.     

Electrotonic transmission between primary afferent fibers and the motor neurons of the isolated spinal cord of various representative amphibians

Studies have been made of monosynaptic excitatory postsynaptic potentials elicited by stimulation of the posterior root in motoneurones of the isolated spinal cord of the frog Rana ridibunda, toad Bufo bufo, and clawed toad Xenopus laevis. In all the amphibians studied, the early component of monosynaptic EPSP was not blocked in Ca-free medium containing 2 mM Mn2+. It is suggested that electrical coupling in anuran amphibians reflects certain stage of evolution of the synaptic transmission in vertebrates.     

Climatic niche conservatism and the evolutionary dynamics in species range boundaries: global congruence across mammals and amphibians

Aim Comparative evidence for phylogenetic niche conservatism – the tendency for lineages to retain their ancestral niches over long time scales – has so far been mixed, depending on spatial and taxonomic scale. We quantify and compare conservatism in the climatic factors defining range boundaries in extant continental mammals and amphibians in order to identify those factors that are most evolutionarily conserved, and thus hypothesized to have played a major role in determining the geographic distributions of many species. We also test whether amphibians show stronger signals of climatic niche conservatism, as expected from their greater physiological sensitivity and lower dispersal abilities. Location Global; continental land masses excluding Antarctica. Methods We used nearly complete global distributional databases to estimate the climatic niche conservatism in extant continental mammals and amphibians. We characterized the climatic niche of each species by using a suite of variables and separately investigate conservatism in each variable using both taxonomic and phylogenetic approaches. Finally, we explored the spatial, taxonomic and phylogenetic patterns in recent climatic niche evolution. Results Amphibians and mammals showed congruent patterns of conservatism in cold tolerance, with assemblages of escapee species (i.e. those escaping most from the climatic constraints of their ancestors) aggregated in the North Temperate Zone. Main conclusions The relative strength of climatic niche conservatism varies across the variables tested, but is strongest for cold tolerance in both mammals and amphibians. Despite the apparent conservatism in this variable, there is also a strong signal of recent evolutionary shifts in cold tolerance in assemblages inhabiting the North Temperate Zone. Our results thus indicate that distribution patterns of both taxa are influenced by both niche conservatism and niche evolution.     

The ascidian mouth opening is derived from the anterior neuropore: Reassessing the mouth/neural tube relationship in chordate evolution

The relative positions of the brain and mouth are of central importance for models of chordate evolution. The dorsal hollow neural tube and the mouth have often been thought of as developmentally distinct structures that may have followed independent evolutionary paths. In most chordates however, including vertebrates and ascidians, the mouth primordia have been shown to fate to the anterior neural boundary. In ascidians such as Ciona there is a particularly intimate relationship between brain and mouth development, with a thin canal connecting the neural tube lumen to the mouth primordium at larval stages. This so-called neurohypophyseal canal was previously thought to be a secondary connection that formed relatively late, after the independent formation of the mouth primordium and the neural tube. Here we show that the Ciona neurohypophyseal canal is present from the end of neurulation and represents the anteriormost neural tube, and that the future mouth opening is actually derived from the anterior neuropore. The mouth thus forms at the anterior midline transition between neural tube and surface ectoderm. In the vertebrate Xenopus, we find that although the mouth primordium is not topologically continuous with the neural tube lumen, it nonetheless forms at this same transition point. This close association between the mouth primordium and the anterior neural tube in both ascidians and amphibians suggests that the evolution of these two structures may be more closely linked than previously appreciated.     

Energy and the tempo of evolution in amphibians

Aim The evolutionary speed hypothesis (ESH) attempts to explain global patterns of species richness on the basis that rates of molecular evolution and speciation in warmer climates have led to a greater accumulation of taxa at lower latitudes. A substantial alternative hypothesis to the ESH is the tropical conservatism hypothesis (TCH). However, recent tests of the TCH, using amphibians as the model taxon, have relied on the assumption that rates of molecular evolution are stable across latitudes and elevations. Here, we test for the first time for systematic variation in rates of molecular evolution across latitude and elevation among amphibians. Location The dataset is geographically diverse with samples from all continents except Antarctica and also from many of the earth's major tropical–warm temperate archipelagos. Methods We tested for substitution rate heterogeneity across climatically varying habitats with the mitochondrial RNA genes 12S and 16S. Thus, we report here on our findings for amphibians – a taxon whose phylogenetic and trophic contexts are remote from those previously tested – using genes that have also not been examined before. The study utilized paired contrasts of sister species (188 species across 18 families, including both caudates and anurans) that are spatially separated in either latitudinal or elevational dimensions. Results We found substantially faster substitution rates for species living in warmer habitats (P= 0.001–0.002) at both lower latitudes (P < 0.02) and lower elevations (P < 0.01). Main conclusions The consistency of these results with the previous studies that used quite different organisms – and in this instance also using different genes – suggests that this is a ubiquitous pattern in nature consistent with the predictions of the ESH. Recent tests of the TCH that, in estimating diversification rates, have relied on the assumption that DNA evolution occurs at a constant rate across latitudes and elevations, require reconsideration in light of the findings presented here. Our results indicate that greater caution is required when estimating dates of divergence using DNA sequence data.