Southern Ocean diversity: new paradigms from molecular ecology

Southern Ocean biodiversity reflects past climate, oceanographic, and tectonic changes. Molecular data from contemporary populations carry signatures of these processes. Here, we review new molecular studies on Southern Ocean benthic fauna. Many of these studies focus on species with extensive geographic or bathymetric distributions, and resolve taxonomic questions. Reviewing all available data, we show that, in addition to reflecting life-history characteristics, the molecular signals found in these studies provide an insight into how species survived the last glacial maximum (LGM). We identify molecular signatures that are characteristic of surviving glacial cycles in small refugia on the continental shelf and distinguish them from molecular signatures that are indicative of surviving glacial cycles in the deep sea.     

The molecular ecology of microbial eukaryotes unveils a hidden world.

In spite of the great success of small-subunit ribosomal RNA (SSU rRNA)-based studies for the analysis of environmental prokaryotic diversity, this molecular approach has seldom been applied to microbial eukaryotes. Recent molecular surveys of the smallest eukaryotic planktonic fractions at different oceanic surface regions and in deep-sea Antarctic samples revealed an astonishing protist diversity. Many of the phylotypes found in the photic region affiliate with photosynthetic groups that are known to contain picoeukaryotic representatives in the range 1-2 microm. Surprisingly, a vast diversity of presumably heterotrophic or mixotrophic lineages is also found. Among these, several novel lineages of heterokonts, and a large diversity of alveolates clustering in two major groups (Groups I and II), are present at all depths in the water column. Many of these new phylotypes appear biogeographically ubiquitous. These initial studies suggest that a wide diversity of small eukaryotes remains to be discovered not only in the ocean but also in other environments. For both ecology and evolutionary studies, it is predicted that environmental molecular identification of eukaryotes will have a profound impact in the immediate future.     

A molecular survey of protist diversity through the central Arctic Ocean

The protist assemblage in the central Arctic Ocean is scarcely surveyed despite them being the major primary producers. Elucidating their response to changing environmental variables requires an a priori analysis of their current diversity, including abundant and rare species. In late summer 2011, samples were collected during the ARK-XXVI/3 expedition (RV Polarstern) to study Arctic protist community structures, by implementation of automated ribosomal intergenic spacer analysis (ARISA) and 454-pyrosequencing. Protist assemblages were related to the hydrology and environmental variables (temperature, salinity, ice coverage, nitrate, phosphate, and silicate). The abundant (≥1 %) biosphere and rare (<1 %) biosphere were considered separately in the diversity analysis in order to reveal their mutual relationships. A relation between hydrology and protist community structure was highly supported by ARISA and partially by 454-pyrosequencing. Sea ice showed a stronger influence on the local community structure than nutrient availability, making statements on the water mass influence more difficult. Dinoflagellates (Syndiniales), chlorophytes (Micromonas spp.), and haptophytes (Phaeocystis spp.) were important contributors to the abundant biosphere, while other dinoflagellates and stramenopiles dominated the rare biosphere. No significant correlation was found between the abundant and rare biosphere. However, relative contributions of major taxonomic groups revealed an unexpected stable community structure within the rare biosphere, indicating a potential constant protist reservoir. This study provides a first molecular survey of protist diversity in the central Arctic Ocean, focusing on the diversity and distribution of abundant and rare protists according to the environmental conditions, and can serve as baseline for future analysis.     

Contrasting seasonal niche separation between rare and abundant taxa conceals the extent of protist diversity

With the advent of molecular methods, it became clear that microbial biodiversity had been vastly underestimated. Since then, species abundance patterns were determined for several environments, but temporal changes in species composition were not studied to the same level of resolution. Using massively parallel sequencing on the 454 GS FLX platform we identified a highly dynamic turnover of the seasonal abundance of protists in the Austrian lake Fuschlsee. We show that seasonal abundance patterns of protists closely match their biogeographic distribution. The stable predominance of few highly abundant taxa, which previously led to the suggestion of a low global protist species richness, is contrasted by a highly dynamic turnover of rare species. We suggest that differential seasonality of rare and abundant protist taxa explains the—so far—conflicting evidence in the ‘everything is everywhere’ dispute. Consequently temporal sampling is basic for adequate diversity and species richness estimates.     

Integrating molecular tools into freshwater ecology: developments and opportunities

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The state of freshwater ecology

1. The case is advanced that freshwater ecologists need to champion the relevance of their work to the development of ecological theory and to the understanding of ecosystem function and behaviour, not least for its importance in addressing pressing applications to the stewardship of the biosphere. An essential step is to review, and update where necessary, the paradigms of aquatic ecology. 2. It is proposed that the major constraint on the organisms, their attributes and adaptations are related first to the physical properties of the medium in which they live. The drives to grow and reproduce relate to the trophic transfer of reduced carbon with important microbial interventions. General principles of ‘emergy’ apply. The supportive capacities of given environments may be set by chemical constraints, but it is suggested that, with the exception of chronically resource-deficient waters, population dynamics relate to opportunities incumbent upon system variability and the consequent pulsation of resources. 3. Variability affects diversity, through frequent revision of the thermodynamic base. Frequent structural change promotes species diversity and, because function is maintained, it appears that efficient function is dependent upon high diversity. Caution is necessary because high productivity and high diversity are both products of the disturbances consequent upon external forcing and manifestly non-equilibrium conditions. 4. Reactions to these statements are canvassed.     

Chromatin domains in higher eukaryotes: insights from genome-wide mapping studies

In genomes of higher eukaryotes, adjacent genes often show coordinated regulation of their expression. Compartmentalization of multiple neighboring genes into a shared chromatin environment can facilitate this coordinated expression. New mapping techniques have begun to reveal that such multigene chromatin domains are a common feature of fly and mammalian genomes. Many different types of chromatin domains have been identified based on the genomic binding patterns of various proteins and histone modifications. In addition, maps of genome–nuclear lamina associations and of looping interactions between loci provide the first systematic views of the three-dimensional folding of interphase chromosomes. These genome-wide datasets uncover new architectural principles of eukaryotic genomes and indicate that multigene chromatin domains are prevalent and important regulatory units.     

Origin of multicellular eukaryotes - insights from proteome comparisons

The complete genomes of the yeast Saccharomyces cerevisiae and the nematode worm Caenorhabditis elegans have recently become available allowing the comparison of the complete protein sets of a unicellular and multicellular eukaryote for the first time. These comparisons reveal some striking trends in terms of expansions or extensive shuffling of specific domains that are involved in regulatory functions and signaling. Similar comparisons with the available sequence data from the plant Arabidopsis thaliana produce consistent results. These observations have provided useful insights regarding the origin of multicellular organisms.     

Evolutionary ecology of learning: insights from fruit flies

Ecologically and evolutionarily oriented research on learning has traditionally been carried out on vertebrates and bees. While less sophisticated than those animals, fruit flies (Drosophila) are capable of several forms of learning, and have the advantage of a short generation time, which makes them an ideal system for experimental evolution studies. This review summarizes the insights into evolutionary questions about learning gained in the last decade from evolutionary experiments on Drosophila. These experiments demonstrate that Drosophila has the genetic potential to evolve a substantially improved learning performance in ecologically relevant learning tasks. In at least one set of selected populations, the improved learning generalized to a task other than that used to impose selection, involving a different behavior, different stimuli, and a different sensory channel for the aversive reinforcement. This improvement in learning ability was associated with reductions in other fitness-related traits, such as larval competitive ability and lifespan, pointing to evolutionary trade-offs for improved learning. These trade-offs were confirmed by other evolutionary experiments where a reduction in learning performance was observed as a correlated response to selection for tolerance to larval nutritional stress or for delayed aging. Such trade-offs could be one reason why fruit flies have not fully used up their evolutionary potential for learning. Finally, another evolutionary experiment with Drosophila provided the first direct evidence for the long-standing idea that learning can under some circumstances accelerate and in others slow down genetically based evolutionary change. These results demonstrate the usefulness of fruit flies as a model system to address evolutionary questions about learning.     

Evolutionary ecology of specialization: insights from phylogenetic analysis

In this Special feature, we assemble studies that illustrate phylogenetic approaches to studying salient questions regarding the effect of specialization on lineage diversification. The studies use an array of techniques involving a wide-ranging collection of biological systems (plants, butterflies, fish and amphibians are all represented). Their results reveal that macroevolutionary examination of specialization provides insight into the patterns of trade-offs in specialized systems; in particular, the genetic mechanisms of trade-offs appear to extend to very different aspects of life history in different groups. In turn, because a species may be a specialist from one perspective and a generalist in others, these trade-offs influence whether we perceive specialization to have effects on the evolutionary success of a lineage when we examine specialization only along a single axis. Finally, how geographical range influences speciation and extinction of specialist lineages remains a question offering much potential for further insight.     

The southwestern Carpathians as an ancient centre of diversity of freshwater gammarid amphipods: insights from the Gammarus fossarum species complex

Gammarus fossarum is a diverse species complex of epigean freshwater amphipods throughout Europe. Due to their poor dispersal capabilities and ubiquity, these crustaceans may serve as a model for investigating the influence of historical factors on the contemporary distribution and diversity patterns of freshwater macrozoobenthos. Here, we investigate the fine‐scale phylogeographic structure of this complex across its range in the southwestern Carpathian Mountains, which comprises two areas that are geographically isolated from its main European distribution area as well as from each other. Given the Tertiary age of many freshwater Gammarus species, we hypothesize that the southwestern Carpathian populations reflect a relict distribution pattern. We used two mitochondrial and three nuclear markers from 32 localities to reconstruct phylogenetic relationships and estimate the timings of divergence among southwestern Carpathian and non‐Carpathian lineages. Cryptic diversity was evaluated from mitochondrial markers by employing phylogenetic and distance‐based methods. We distinguished at least 16 cryptic microendemic taxa, some of them coexisting, distributed in the southwestern Carpathians in a mosaic‐like pattern. These lineages form a monophyletic group together with several lineages from southeastern Europe. Estimated divergence times indicate a Middle Miocene origin of this clade, with many deep splits dating back to more than 10 Ma. This time frame corresponds with a period of intense geological subsidence in the region that gave birth to the Pannonian Basin. We conclude that subsidence could have been an important driver of diversification in freshwater Gammarus and that the southwestern Carpathians represent an ancient centre of diversity for these crustaceans.     

Impact of ice ages on circumpolar molecular diversity: insights from an ecological key species

We address the impact of the ice age cycles on intraspecific cpDNA diversity, for the first time on the full circumboreal-circumarctic scale. The bird-dispersed bog bilberry (or arctic blueberry, Vaccinium uliginosum) is a key component of northern ecosystems and is here used to assess diversity in previously glaciated vs. unglaciated areas and the importance of Beringia as a refugium and source for interglacial expansion. Eighteen chloroplast DNA haplotypes were observed in and among 122 populations, grouping into three main lineages which probably diverged before, and thus were affected more or less independently by, all major glaciations. The boreal 'Amphi-Atlantic lineage' included one haplotype occurring throughout northern Europe and one occurring in eastern North America, suggesting expansion from at least two bottlenecked, glacial refugium populations. The boreal 'Beringian lineage' included seven haplotypes restricted to Beringia and the Pacific coast of USA. The 'Arctic-Alpine lineage' included nine haplotypes, one of them fully circumpolar. This lineage was unexpectedly diverse, also in previously glaciated areas, suggesting that it thrived on the vast tundras during the ice ages and recolonized deglaciated terrain over long distances. Its largest area of persistence during glaciations was probably situated in the north, stretching from Beringia and far into Eurasia, and it probably also survived the last glaciation in southern mountain ranges. Although Beringia apparently was important for the initial divergence and expansion of V. uliginosum as well as for continuous survival of both the Beringian and Arctic-Alpine lineages during all ice ages, this region played a minor role as a source for later interglacial expansions.     

Analyses of genetic ancestry enable key insights for molecular ecology

Gene flow and recombination in admixed populations produce genomes that are mosaic combinations of chromosome segments inherited from different source populations, that is, chromosome segments with different genetic ancestries. The statistical problem of estimating genetic ancestry from DNA sequence data has been widely studied, and analyses of genetic ancestry have facilitated research in molecular ecology and ecological genetics. In this review, we describe and compare different model‐based statistical methods used to infer genetic ancestry. We describe the conceptual and mathematical structure of these models and highlight some of their key differences and shared features. We then discuss recent empirical studies that use estimates of genetic ancestry to analyse population histories, the nature and genetic basis of species boundaries, and the genetic architecture of traits. These diverse studies demonstrate the breadth of applications that rely on genetic ancestry estimates and typify the genomics‐enabled research that is becoming increasingly common in molecular ecology. We conclude by identifying key research areas where future studies might further advance this field.     

SppA peptidases: family diversity from heterotrophic bacteria to photoautotrophic eukaryotes

Serine-type SppA peptidases are found in viruses, archaea, eubacteria and in chloroplasts. The family consists of two homologous groups of proteins, designated SppA1 and SppA2, which differ in molecular weight and domain organization. SppA1 is a high molecular weight protein of 70 kDa with two homologous domains with protease signatures. SppA2 represents a half-size SppA1 comprised of only one protease domain. The genomes of most heterotrophic and photosynthetic bacteria encode both, SppA1 and SppA2 proteins, while genome of higher plants, Arabidopsis and rice, encode only one protease, SppA1. An intriguing feature of SppA proteases is their similarity to ClpP proteases in catalytically active regions. SppA functions and gene expression differ between heterotrophic and photosynthetic organisms, in which SppA expression is light-regulated. Apparently, SppA regulation and substrate specificities were modified in photosynthetic organisms and are essential for acclimation of the thylakoid membrane system to light.     

Lateral transfer of eukaryotic ribosomal RNA genes: an emerging concern for molecular ecology of microbial eukaryotes

Ribosomal RNA (rRNA) genes are widely utilized in depicting organismal diversity and distribution in a wide range of environments. Although a few cases of lateral transfer of rRNA genes between closely related prokaryotes have been reported, it remains to be reported from eukaryotes. Here, we report the first case of lateral transfer of eukaryotic rRNA genes. Two distinct sequences of the 18S rRNA gene were detected from a clonal culture of the stramenopile, Ciliophrys infusionum. One was clearly derived from Ciliophrys, but the other gene originated from a perkinsid alveolate. Genome-walking analyses revealed that this alveolate-type rRNA gene is immediately adjacent to two protein-coding genes (ubc12 and usp39), and the origin of both genes was shown to be a stramenopile (that is, Ciliophrys) in our phylogenetic analyses. These findings indicate that the alveolate-type rRNA gene is encoded on the Ciliophrys genome and that eukaryotic rRNA genes can be transferred laterally.     

土壤真菌多样性及分子生态学研究进展

真菌是土壤中一类重要的微生物,参与有机质分解,与植物共生为植物提供养分,同时病原真菌也会引起粮食产量的降低．土壤真菌多样性在维持生态系统的平衡和为人类提供大量未开发资源上起到了独特而重要的作用．本文从物种多样性、生境多样性、功能多样性角度阐述了土壤真菌多样性,并从农田、林地、草地、极端环境与一些复杂环境土壤真菌多样性层面综述了土壤真菌多样性分子生态学研究进展。同时论述了一些影响真菌多样性的因素,并对土壤真菌多样性研究的前景提出展望．     

The emergence and maintenance of diversity: insights from experimental bacterial populations

Mechanisms maintaining genetic and phenotypic variation in natural populations are central issues in ecology and evolution. However, the long generation times of most organisms and the complexity of natural environments have made elucidation of ecological and evolutionary mechanisms difficult. Experiments using bacterial populations propagated in controlled environments reduce ecosystem complexity to the point where understanding simple processes in isolation becomes possible. Recent studies reveal the circumstances and mechanisms that promote the emergence of stable polymorphisms.     

The effects of pollen diversity on plant reproduction: insights from apple

For many plants, the number of pollen genotypes deposited on a flower’s stigma is positively related to fruit maturation and seed number; however, the mechanisms underlying this effect are less understood. Here we examined whether diversity of pollen (1, 3, or 5 donors) affects reproductive success in self-incompatible apple (Malus × domestica). Using paternity analysis, we then assessed the siring rate of individual donors to test whether diversity effects are related to the presence of a superior pollen donor or to donor × donor interactions. Increasing the diversity of compatible pollen enhanced seed number and reduced seed abortion in some but not all recipient genotypes. This effect was associated with an increased number of sires per fruit and non-random siring success among pollen donors. Two donors had consistently high siring rates, regardless of recipient genotype; however, the siring success of pollen donors was not correlated with their siring success in single-donor pollinations. Rather, siring success was affected by the identity of other pollen genotypes present on the stigma. Our results therefore suggest that the effects of pollen diversity are variable but may enhance fecundity by fostering interactions between pollen donors.