Hsp90 in non-mammalian metazoan model systems

The molecular chaperone Hsp90 has been discovered in the heat-shock response of the fruit fly more than 30years ago. Today, it is becoming clear that Hsp90 is in the middle of a regulatory system, participating in the modulation of many essential client proteins and signaling pathways. Exerting these activities, Hsp90 works together with about a dozen of cochaperones. Due to their organismal simplicity and the possibility to influence their genetics on a large scale, many studies have addressed the function of Hsp90 in several multicellular model systems. Defined pathways involving Hsp90 client proteins have been identified in the metazoan model systems of Caenorhabditis elegans, Drosophila melanogaster and the zebrafish Danio rerio. Here, we summarize the functions of Hsp90 during muscle maintenance, development of phenotypic traits and the involvement of Hsp90 in stress responses, all of which were largely uncovered using the model organisms covered in this review. These findings highlight the many specific and general actions of the Hsp90 chaperone machinery. This article is part of a Special Issue entitled: Heat Shock Protein 90 (HSP90).     

基部藓类分子系统学研究

基部藓类是稳定地处于藓类系统发育树基部的类群.它包括7纲,2亚纲,10目,10科,34属,637种.基部藓类虽然只占藓类种类的5％,但由于其内部各类群孢子体形态极为丰富,因此对于理解整个藓类植物的系统发育具有重要意义.通过对48个种(36个藓类、4个维管植物、2个角苔、4个苔类和2个藻类)的9个DNA片段(NU:26S,18S; MT:nad5,cox1; CP:rbcL,rps4,cp-LSU,cp-SSU,atpB)进行分子系统学分析,综合最大似然法(maximum likelihood)、最大简约法(most parsimony)和贝叶斯分析(Bayesian inference)方法的建树结果,理清了前人研究中存在冲突的类群之间的关系并为已确定的关系提供了更高的支持率.研究结果如下:(1)藻苔纲和泥炭藓纲互为姐妹类群,处于整个藓类的最基部;(2)黑藓纲与黑真藓纲互为姐妹类群(3)长台藓纲和具齿藓类组成单系;(4)四齿藓纲是所有具齿藓类的基部类群;(5)烟杆藓亚纲处于真藓纲的最基部,其次是短颈藓亚纲.以上结论在分子系统树上得到了很高的支持率.     

The boundaries of river systems: the metazoan perspective

1. This overview of metazoans associated with the riparian/groundwater interface focuses on the fauna inhabiting substratum interstices within the stream bed and in alluvial aquifers beneath the floodplain. The objective is to integrate knowledge of habitat conditions and ecology of the interstitial fauna into a broad spatiotemporal perspective of lotic ecosystems. 2. Most aquatic metazoans of terrestrial ancestry, secondarily aquatic forms including insects and water mites (Hydracarina), are largely confined to surface waters (epigean), most of the time penetrating only the superficial interstices of the stream bed. 3. Primary aquatic metazoans include crustaceans and other groups whose entire evolutionary histories took place in water. Some species are epigean, whereas other members of the primary aquatic fauna are true subterranean forms (hypogean ) , residing deep within the stream bed and in alluvial aquifers some distance laterally from the channel. 4. The hypogean/epigean affinities of interstitial animals are reflected in repetitive gradients of species distribution patterns along vertical (depth within the stream bed), longitudinal (riffle/pool), and lateral (across the floodplain) spatial dimensions, as well as along recovery trajectories following floods (temporal dimension). 5. Fluvial dynamics and sediment characteristics interact to determine hydraulic conductivity, oxygen levels, pore space, particle size heterogeneity, organic content and other habitat conditions within the interstitial milieu. 6. Multidimensional environmental gradients occur at various scales across riparian/groundwater boundary zones. The spatiotemporal variability of hydrogeomorphological processes plays an important role in determining habitat heterogeneity, habitat stability, and connectivity between habitat patches, thereby structuring biodiversity patterns across the riverine landscape. 7. The erosive action of flooding maintains a diversity of hydrarch and riparian successional stages in alluvial floodplains. The patchy distribution patterns of interstitial communities at the floodplain scale reflect, in part, the spatial heterogeneity engendered by successional processes. 8. Interstitial metazoans engage in passive and active movements between surface waters and ground waters, between aquatic and riparian habitats, and between different habitat types within the lotic system. Some of these are extensive migrations that involve significant exchange of organic matter and energy between ecosystem compartments. 9. The generally high resilience of lotic ecosystems to disturbance is attributable, in part, to high spatiotemporal heterogeneity. Habitat patches less affected by a particular perturbation may serve as ’refugia ‘; from which survivors recolonize more severely affected areas. Mechanisms of refugium use may also occur within habitats, as, for example, through ontogenetic shifts in microhabitat use. Rigorous investigations of interstitial habitats as refugia should lead to a clearer understanding of the roles of disturbance and stochasticity in lotic ecosystems. 10. Development of realistic ’whole river ‘; food webs have been constrained by the exclusion of interstitial metazoans, which may in fact contribute the majority of energy flow in lotic ecosystems. A related problem is failure to include groundwater/riparian habitats as integral components of alluvial rivers. A conceptual model is presented that integrates groundwater and riparian systems into riverine food webs and that reflects the spatiotemporal complexity of the physical system and connectivity between different components. 11. Interstitial metazoans also serve as ’ecosystem engineers, ‘; by influencing the availability of resouces to other species and by modifying habitat conditions within the sediment. For example, by grazing on biofilm, interstitial animals may markedly stimulate bacterial growth rates and nutrient dynamics. 12. Although there has been a recent surge of interest in the role of interstitial animals in running waters, the knowledge gaps are vast. For example, basic environmental requirements of the majority of groundwater metazoans remain uninvestigated. Virtually nothing is known regarding the role of biotic interactions in structuring faunal distribution patterns across groundwater/riparian boundary zones. Interstitial metazoans may contribute significantly to the total productivity and energy flow of the biosphere, but such data are not available. Nor are sufficient data available to determine the contribution of groundwater animals to estimates of global biodiversity. 13. Effective ecosystem management must include groundwater/riparian ecotones and interstitial metazoans in monitoring and restoration efforts. Evidence suggests that a ’connected ‘; groundwater/riparian system provides natural pollution control, prevents clogging of sediment interstices and maintains high levels of habitat heterogeneity and successional stage diversity. River protection and restoration should maintain or re-establish at least a portion of the natural fluvial dynamics that sustains the ecological integrity of the entire riverine–floodplain–aquifer ecosystem. Keywords: groundwater/riparian ecotones, hyporheic habitat, epigean, hypogean, interstitial fauna, biodiversity, food webs     

New Early Eocene Basal tapiromorph from Southern China and Its Phylogenetic Implications

A new Early Eocene tapiromorph, Meridiolophus expansus gen. et sp. nov., from the Sanshui Basin, Guangdong Province, China, is described and discussed. It is the first reported Eocene mammal from the basin. The new taxon, represented by a left fragmentary mandible, is characterized by an expanded anterior symphyseal region, a long diastema between c1 and p1, a rather short diastema between p1 and p2, smaller premolars relative to molars, an incipient metaconid appressed to the protoconid on p3, a prominent entoconid on p4, molar metaconid not twinned, cristid obliqua extending mesially and slightly lingually from the hypoconid, inclined metalophid and hypolophid, and small hypoconulid on the lower preultimate molars. Meridiolophus is morphologically intermediate between basal Homogalax-like taxa and derived tapiromorphs (such as Heptodon). Phylogenetic analysis indicates Equidae is more closely related to Tapiromorpha than to Palaeotheriidae, although the latter is only represented by a single species Pachynolophus eulaliensis. ‘Isectolophidae’, with exception of Meridiolophus and Karagalax, has the closest affinity with Chalicotherioidea. Furthermore, the majority rule consensus tree shows that Meridiolophus is closer to Karagalax than to any other ‘isectolophid’, and both genera represent stem taxa to crown group Ceratomorpha.     

Molecular Dissection of the Basal Clades in the Human Y Chromosome Phylogenetic Tree

One hundred and forty-six previously detected mutations were more precisely positioned in the human Y chromosome phylogeny by the analysis of 51 representative Y chromosome haplogroups and the use of 59 mutations from literature. Twenty-two new mutations were also described and incorporated in the revised phylogeny. This analysis made it possible to identify new haplogroups and to resolve a deep trifurcation within haplogroup B2. Our data provide a highly resolved branching in the African-specific portion of the Y tree and support the hypothesis of an origin in the north-western quadrant of the African continent for the human MSY diversity.     

Phylogenetic models and model selection for noncoding DNA

Awareness of the complex structure and evolutionary dynamics of noncoding DNA has improved both noncoding sequence alignment and the use of microstructural changes as characters in phylogenetic analysis. The next step is to consider improvements in the use and selection of phylogenetic models for noncoding sequence data. Models of character evolution are central to phylogeny estimation, but the use of an inadequate model can mislead topology selection and branch length estimations. This is particularly likely when sequence divergence is either limited (nearly invariable, as in population-level or species-level studies) or extreme (nearly saturated, as in deep-level studies that focus on conserved secondary structures). Noncoding data sets are often at these extremes, and they can be particularly awkward for model definition and model selection. This paper introduces the goals of model use in phylogenetics and identifies ten issues that arise from the application of models to noncoding sequence data. It is concluded that most of these issues derive from small data set sizes, very low or very high sequence variability, limitations of current phylogenetic models, and possibly character definition and nonindependence. Recommendations are made that should help to improve alignment, character quality, model selection, and phylogeny estimation based on noncoding sequence data.     

Phylogenetic inference under the pure drift model

When pairwise genetic distances are used for phylogenetic reconstruction, it is usually assumed that the genetic distance between two taxa contains information about the time after the two taxa diverged. As a result, upon an appropriate transformation if necessary, the distance usually can be fitted to a linear model such that it is expressed as the sum of lengths of all branches that connect the two taxa in a given phylogeny. This kind of distance is referred to as "additive distance." For a phylogenetic tree exclusively driven by random genetic drift, genetic distances related to coancestry coefficients (theta XY) between any two taxa are more suitable. However, these distances are fundamentally different from the additive distance in that coancestry does not contain any information about the time after two taxa split from a common ancestral population; instead, it reflects the time before the two taxa diverged. In other words, the magnitude of theta XY provides information about how long the two taxa share the same evolutionary pathways. The fundamental difference between the two kinds of distances has led to a different algorithm of evaluating phylogenetic trees when theta XY and related distance measures are used. Here we present the new algorithm using the ordinary- least-squares approach but fitting to a different linear model. This treatment allows genetic variation within a taxon to be included in the model. Monte Carlo simulation for a rooted phylogeny of four taxa has verified the efficacy and consistency of the new method. Application of the method to human population was demonstrated.      

The replicon revisited: an old model learns new tricks in metazoan chromosomes

The origins of DNA replication were proposed in the replicon model to be specified genetically by replicator elements that coordinate the initiation of DNA synthesis with gene expression and cell growth. Recent studies have identified DNA sequences in mammalian cells that fulfil the genetic criteria for replicators and are beginning to uncover the sequence requirements for the initiation of DNA replication. Mammalian replicators are com- posed of non-redundant modules that cooperate to direct initiation to specific chromosomal sites. Conversely, replicators do not show strong sequence similarity, and their ability to initiate replication depends on the chromosomal context and epigenetic factors, as well as their primary sequence. Here, we review the properties of metazoan replicators, and discuss the genetic and epigenetic factors that determine where and when DNA replication is initiated.     

Social context predicts recognition systems in ant queens

Recognition of group‐members is a key feature of sociality. Ants use chemical communication to discriminate nestmates from intruders, enhancing kin cooperation and preventing parasitism. The recognition code is embedded in their cuticular chemical profile, which typically varies between colonies. We predicted that ants might be capable of accurate recognition in unusual situations when few individuals interact repeatedly, as new colonies started by two to three queens. Individual recognition would be favoured by selection when queens establish dominance hierarchies, because repeated fights for dominance are costly; but it would not evolve in absence of hierarchies. We previously showed that Pachycondyla co‐founding queens, which form dominance hierarchies, have accurate individual recognition based on chemical cues. Here, we used the ant Lasius niger to test the null hypothesis that individual recognition does not occur when co‐founding queens do not establish dominance hierarchies. Indeed, L. niger queens show a similar level of aggression towards both co‐foundresses and intruders, indicating that they are unable of individual recognition, contrary to Pachycondyla. Additionally, the variation in chemical profiles of Lasius and Pachycondyla queens is comparable, thus informational constraints are unlikely to apply. We conclude that selection pressure from the social context is of crucial significance for the sophistication of recognition systems.     

Technological advances and genomics in metazoan parasites

Molecular biology has provided the means to identify parasite proteins, to define their function, patterns of expression and the means to produce them in quantity for subsequent functional analyses. Whole genome and expressed sequence tag programmes, and the parallel development of powerful bioinformatics tools, allow the execution of genome-wide between stage or species comparisons and meaningful gene-expression profiling. The latter can be undertaken with several new technologies such as DNA microarray and serial analysis of gene expression. Proteome analysis has come to the fore in recent years providing a crucial link between the gene and its protein product. RNA interference and ballistic gene transfer are exciting developments which can provide the means to precisely define the function of individual genes and, of importance in devising novel parasite control strategies, the effect that gene knockdown will have on parasite survival.     

Near intron pairs and the metazoan tree

Gene structure data can substantially advance our understanding of metazoan evolution and deliver an independent approach to resolve conflicts among existing hypotheses. Here, we used changes of spliceosomal intron positions as novel phylogenetic marker to reconstruct the animal tree. This kind of data is inferred from orthologous genes containing mutually exclusive introns at pairs of sequence positions in close proximity, so-called near intron pairs (NIPs). NIP data were collected for 48 species and utilized as binary genome-level characters in maximum parsimony (MP) analyses to reconstruct deep metazoan phylogeny. All groupings that were obtained with more than 80% bootstrap support are consistent with currently supported phylogenetic hypotheses. This includes monophyletic Chordata, Vertebrata, Nematoda, Platyhelminthes and Trochozoa. Several other clades such as Deuterostomia, Protostomia, Arthropoda, Ecdysozoa, Spiralia, and Eumetazoa, however, failed to be recovered due to a few problematic taxa such as the mite Ixodes and the warty comb jelly Mnemiopsis. The corresponding unexpected branchings can be explained by the paucity of synapomorphic changes of intron positions shared between some genomes, by the sensitivity of MP analyses to long-branch attraction (LBA), and by the very unequal evolutionary rates of intron loss and intron gain during evolution of the different subclades of metazoans. In addition, we obtained an assemblage of Cnidaria, Porifera, and Placozoa as sister group of Bilateria + Ctenophora with medium support, a disputable, but remarkable result. We conclude that NIPs can be used as phylogenetic characters also within a broader phylogenetic context, given that they have emerged regularly during evolution irrespective of the large variation of intron density across metazoan genomes.     

Phylogenetic reconstruction of parental-care systems in the ancestors of birds

Due to the controversy surrounding incipient avian parental care, ancestral parental care systems were reconstructed in a phylogeny including major extant amniote lineages. Using two different resolutions for the basal avian branches, transitions between the states no care, female care, biparental care and male care were inferred for the most basal branches of the tree. Uniparental female care was inferred for the lineage to birds and crocodiles. Using a phylogeny where ratites and tinamous branch off early and an ordered character-state assumption, a transition to biparental care was inferred for the ancestor of birds. This ancestor could be any organism along the lineage leading from the crocodile-bird split up to modern birds, not necessarily the original bird. We discuss the support for alternative avian phylogenies and the homology in parental care between crocodiles and birds. We suggest that the phylogenetic pattern should be used as a starting point for a more detailed analysis of parental care systems in birds and their relatives.     

Industrial bioprocess control and optimization in the context of systems biotechnology

The developments of the systems biotechnology and its application in the industrial process open up new horizons to industrial biotechnology. The unprecedented understanding of the relationships between cellular behaviors and the surrounding environments during the bioprocess has been achieved. In this paper, we review new advances in the strain improvement, bioprocess control and optimization. The holistic viewpoints and ideas applied in industrial bioprocesses and their future prospects are discussed by illustrating some successful cases.     

Combinatorial model for the formation of body plans in higher metazoan taxa: Paleontological insight

The body plans of higher metazoan taxa were formed during a short time (on the geological time scale) by combination of the previously developed characters. The combinations were realized as a result of manifestation of latent characters in adults through various heterochronies. This resulted in mosaic evolution and concealment of intermediate forms. Many characters of new body plans appeared in the ancestral taxon and their various combinations in the newly established taxa formed the archaic diversity. The maximum rank of newly appearing higher taxa decreased with geological time. The evolution of metazoans passed from the development of the general body plan to less significant details and appearance of body plans describing taxa of lower ranks. New body plans of higher taxa were superposed on the old body plan rather than replaced it, extending with time the subordination of body plans and respective hierarchy of taxa. Aromorphoses are always connected with the appearance of a new body plan. The appearance of new taxa and an increase in morphological diversity mostly occurred at certain boundaries in the development of the biota, which were connected with a sharp increase in the previously limited resources.