南极冰下水生态系统微生物与生源元素循环研究进展

南极大陆冰盖下存在液态水,形成了由冰下湖、冰下河/溪、冰封湖和冰架下水体等组成的冰下水生态系统,具有低温、黑暗和寡营养等极端的环境条件特征。微生物主导了南极冰下水生态系统,其具有丰富多样的种群构成、功能形式和独特的适应机制,在生源元素生物地球化学循环过程中起了重要作用。研究南极冰下微生物群落的生态特征及其参与的生源元素地球化学循环过程,可为揭示地球生命演化和探索外星生命提供指示,具有重要的科学意义。本文综述了南极冰下水生态系统的极端环境条件、冰下微生物的多样性、冰下微生物参与的生物地球化学循环以及冰下微生物的适极机理,最后基于研究现状展望了南极冰下微生物的未来研究方向。     

Mercury (micro)biogeochemistry in polar environments

The contamination of polar regions with mercury that is transported as inorganic mercury from lower latitudes has resulted in the accumulation of methylmercury in the food chain of polar environments, risking the health of humans and wildlife. This problem is likely to be particularly severe in coastal marine environments where active cycling occurs. Little is currently known about how mercury is methylated in polar environments. Relating observations on mercury deposition and transport through polar regions to knowledge of the microbiology of cold environments and considering the principles of mercury transformations as have been elucidated in temperate aquatic environments, we propose that in polar regions (1) variable pathways for mercury methylation may exist, (2) mercury bioavailability to microbial transformations may be enhanced, and (3) microbial niches within sea ice are sites where active microorganisms are localized in proximity to high concentrations of mercury. Thus, microbial transformations, and consequently mercury biogeochemistry, in the Arctic and Antarctic are both unique and common to these processes in lower latitudes, and understanding their dynamics is needed for the management of mercury-contaminated polar environments.     

Divergences in the Response to Ultraviolet Radiation Between Polar and Non-Polar Ciliated Protozoa

Ultraviolet (UV) radiation has detrimental effects on marine ecosystems, in particular in the polar regions where stratospheric ozone reduction causes higher levels of solar radiation. We analyzed two polar species of Euplotes, Euplotes focardii and Euplotes nobilii, for the sensitivity to UV radiation in comparison with two akin species from mid-latitude and tropical waters. Results showed that they face UV radiation much more efficiently than the non-polar species by adopting alternative strategies that most likely reflect different times of colonization of the polar waters. While E. focardii, which is endemic to the Antarctic, survives for longer exposed to UV radiation, E. nobilii, which inhabits both the Antarctic and Arctic, recovers faster from UV-induced damage.     

Glaciers and ice sheets as a biome

The tundra is the coldest biome described in typical geography and biology textbooks. Within the cryosphere, there are large expanses of ice in the Antarctic, Arctic and alpine regions that are not regarded as being part of any biome. During the summer, there is significant melt on the surface of glaciers, ice caps and ice shelves, at which point microbial communities become active and play an important role in the cycling of carbon and other elements within the cryosphere. In this review, we suggest that it is time to recognise the cryosphere as one of the biomes of Earth. The cryospheric biome encompasses extreme environments and is typified by truncated food webs dominated by viruses, bacteria, protozoa and algae with distinct biogeographical structures.     

Diel vertical migration of Arctic zooplankton during the polar night

High-latitude environments show extreme seasonal variation in physical and biological variables. The classic paradigm of Arctic marine ecosystems holds that most biological processes slow down or cease during the polar night. One key process that is generally assumed to cease during winter is diel vertical migration (DVM) of zooplankton. DVM constitutes the largest synchronized movement of biomass on the planet, and is of paramount importance for marine ecosystem function and carbon cycling. Here we present acoustic data that demonstrate a synchronized DVM behaviour of zooplankton that continues throughout the Arctic winter, in both open and ice-covered waters. We argue that even during the polar night, DVM is regulated by diel variations in solar and lunar illumination, which are at intensities far below the threshold of human perception. We also demonstrate that winter DVM is stronger in open waters compared with ice-covered waters. This suggests that the biologically mediated vertical flux of carbon will increase if there is a continued retreat of the Arctic winter sea ice cover.     

高寒生态系统微生物群落研究进展

高寒生态系统分布在高纬度或高海拔、气候寒冷的地区,包括北极苔原、高山苔原、青藏高原等.高寒生态系统对气候变化非常敏感,其土壤中储存大量的有机碳,对全球的碳平衡起关键作用.微生物是生物地球化学循环的主要驱动者,微生物群落对气候变化的响应和反馈影响生态系统的功能与稳定性.本文回顾了高寒生态系统微生物群落组成、多样性与空间分布,以及微生物群落对气候变化(增温、氮沉降、火干扰)的响应,为拓展我国高寒生态系统微生物研究提供基础.     

Diverse viromes in polar regions: A retrospective study of metagenomic data from Antarctic animal feces and Arctic frozen soil in 2012–2014

Antarctica and the Arctic are the coldest places, containing a high diversity of microorganisms, including viruses, which are important components of polar ecosystems. However, owing to the difficulties in obtaining access to animal and environmental samples, the current knowledge of viromes in polar regions is still limited. To better understand polar viromes, this study performed a retrospective analysis using metagenomic sequencing data of animal feces from Antarctica and frozen soil from the Arctic collected during 2012–2014. The results reveal diverse communities of DNA and RNA viruses from at least 23 families from Antarctic animal feces and 16 families from Arctic soils. Although the viral communities from Antarctica and the Arctic show a large diversity, they have genetic similarities with known viruses from different ecosystems and organisms with similar viral proteins. Phylogenetic analysis of Microviridae, Parvoviridae, and Larvidaviridae was further performed, and complete genomic sequences of two novel circular replication-associated protein (rep)-encoding single-stranded (CRESS) DNA viruses closely related to Circoviridae were identified. These results reveal the high diversity, complexity, and novelty of viral communities from polar regions, and suggested the genetic similarity and functional correlations of viromes between the Antarctica and Arctic. Variations in viral families in Arctic soils, Arctic freshwater, and Antarctic soils are discussed. These findings improve our understanding of polar viromes and suggest the importance of performing follow-up in-depth investigations of animal and environmental samples from Antarctica and the Arctic, which would reveal the substantial role of these viruses in the global viral community.     

Did glacial advances during the Pleistocene influence differently the demographic histories of benthic and pelagic Antarctic shelf fishes? – Inferences from intraspecific mitochondrial and nuclear DNA sequence diversity

Background  

Circum-Antarctic waters harbour a rare example of a marine species flock – the Notothenioid fish, most species of which are restricted to the continental shelf. It remains an open question as to how they survived Pleistocene climatic fluctuations characterised by repeated advances of continental glaciers as far as the shelf break that probably resulted in a loss of habitat for benthic organisms. Pelagic ecosystems, on the other hand, might have flourished during glacial maxima due to the northward expansion of Antarctic polar waters. In order to better understand the role of ecological traits in Quaternary climatic fluctuations, we performed demographic analyses of populations of four fish species from the tribe Trematominae, including both fully benthic and pelagic species using the mitochondrial cytochrome b gene and an intron from the nuclear S7 gene.     

The influence of nematodes on below-ground processes in grassland ecosystems

This review summarises recent information on beneficial roles that soil nematodes play in the cycling of carbon and other plant nutrients in grassland ecosystems. In particular, we focus on the role of the two dominant functional groups of nematodes, namely the microbial- and root-feeders, and how their activities may enhance soil ecosystem-level processes of nutrient cycling and, ultimately, plant productivity in managed and unmanaged grassland ecosystems. We report recent experiments which show that low amounts of root herbivory by nematodes can increase the allocation of photoassimilate carbon to roots, leading to increased root exudation and microbial activity in the rhizosphere. The effects of these interactions on soil nutrient cycling and plant productivity are discussed. Evidence is presented to show that the feeding activities of microbial-feeding nematodes can enhance nutrient mineralization and plant nutrient uptake in grasslands, but that these responses are highly species-specific and appear to be strongly regulated by higher trophic groups of fauna (top-down regulation). We recommend that future studies of the roles of nematodes in grasslands ecosystems should consider these more complex trophic interactions and also the effects of species diversity of nematodes on soil ecosystem-level processes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Distinct or similar? Soft bottom polychaete diversity in Arctic and Antarctic glacial fjords

The main aim of this study was to compare the polychaete communities in two similar polar areas: an Arctic fjord, Hornsund (Svalbard) and an Antarctic fjord, Ezcurra Inlet (South Shetlands). This is the first attempt to compare Arctic and Antarctic diversity based on fully comparable datasets. Forty van Veen grab samples were collected in each fjord: twenty replicates were taken in each of two fjord areas characterized by a different level of glacial disturbance—in the inner (glacial bay) and outer (fjord mouth) region of both fjords, from depths of about 100 m in 2005 (Hornsund) and in 2007 (Ezcurra Inlet). In the glacial bays, species richness and diversity were significantly higher in Ezcurra Inlet than in Hornsund due to higher rate of glacial disturbance in the latter one. In the outer areas, species richness was similar in both fjords, although diversity values were higher in Ezcurra Inlet. Polychaete species richness in the habitats characterized by similar level of disturbance (outer areas of the fjords) was the same in both polar regions. At this small scale, where community drivers are very similar, the species richness seems to be independent from the local or regional species pool.     

Molecular evolution of haemoglobins of polar fishes

The Arctic and the Antarctic differ by age and isolation of the respective marine faunas. Antarctic fish are highly stenothermal, in response to stable water temperatures, whereas the Arctic ones are exposed to seasonal and latitudinal temperature variations. The knowledge of the mechanisms of phenotypic response to cold exposure in species of both polar habitats offers fundamental insights into the nature of environmental adaptation. In the process of cold adaptation, the evolutionary trend of Antarctic fish has led to unique specialisations, including modification of haematological characteristics, e.g. decreased amounts and multiplicity of haemoglobins.Unlike Antarctic Notothenioidei, Arctic teleosts have high haemoglobin multiplicity. Although the presence of functionally and structurally distinct haemoglobins is a plesiomorphic condition for many perciform-like fishes, it seems that the oxygen-transport system of teleost fish in the Arctic region has been adjusted to temperature differences and fluctuations of Arctic waters, much larger than in the Antarctic. The amino-acid sequences used to gain insight into the evolution history of α and β globins of polar fish have clearly shown that Antarctic and Arctic globins have different phylogenies, leading to the hypothesis that the selective pressure of environment stability allows the phylogenetic signal to be maintained in the Antarctic sequences, whereas environmental variability would tend to disrupt this signal in Arctic sequences.     

The future of soil invertebrate communities in polar regions: different climate change responses in the Arctic and Antarctic?

The polar regions are experiencing rapid climate change with implications for terrestrial ecosystems. Here, despite limited knowledge, we make some early predictions on soil invertebrate community responses to predicted twenty‐first century climate change. Geographic and environmental differences suggest that climate change responses will differ between the Arctic and Antarctic. We predict significant, but different, belowground community changes in both regions. This change will be driven mainly by vegetation type changes in the Arctic, while communities in Antarctica will respond to climate amelioration directly and indirectly through changes in microbial community composition and activity, and the development of, and/or changes in, plant communities. Climate amelioration is likely to allow a greater influx of non‐native species into both the Arctic and Antarctic promoting landscape scale biodiversity change. Non‐native competitive species could, however, have negative effects on local biodiversity particularly in the Arctic where the communities are already species rich. Species ranges will shift in both areas as the climate changes potentially posing a problem for endemic species in the Arctic where options for northward migration are limited. Greater soil biotic activity may move the Arctic towards a trajectory of being a substantial carbon source, while Antarctica could become a carbon sink.     

草藻型稳态转换对湖泊微生物结构及其碳循环功能的影响

湖泊是地球表层系统中水、土、气等各个圈层相互作用的联结点,对区域物质如碳等元素循环具有重要影响.微生物是湖泊等水生态系统中的重要组成部分,是湖泊等生态系统中碳等元素物质循环的主要驱动者,是深入了解湖泊碳循环过程的关键.受人类活动等影响,湖泊生态系统,尤其是浅水湖泊生态系统往往表现出以高等水生植物(草型)为主要初级生产者的清水稳定态和以浮游藻类(藻型)为主要初级生产者的浊水稳定态,而随着湖泊营养负荷和湖泊环境条件的变化,这两个不同的稳定态之间可以发生转换或者剧变,这种剧变不仅影响湖泊生态系统中的微生物结构,而且对湖泊中有机碳的形成、循环过程及其微生物驱动机制产生重大影响.本文重点就湖泊生态系统中有机碳的转换与微生物关系以及草藻型稳定态的转换对微生物结构及其碳循环功能的影响等进行综述,进一步分析其中的关键科学问题,以期为深入了解湖泊生态系统中碳等元素循环的微生物驱动过程与机制提供帮助.     

Unusual fungal niches

Fungi are found in all aerobic ecosystems, colonizing a diversity of substrates and performing a wide diversity of functions, some of which are not well understood. Many spices of fungi are cosmopolitan and generalists or habitats. Unusual fungal niches are habitats where extreme conditions would be expected to prevent the development of a mycobiota. In this review we describe five unusual fungal habitats in which fungi occupy poorly understood niches: Antarctic dry valleys, high Arctic glaciers, salt flats and salterns, hypersaline microbial mats and plant trichomes. Yeasts, black yeast-like fungi, melanized filamentous species as well as representatives of Aspergillus and Penicillium seem to be dominant among the mycobiota adapted to cold and saline niches. Plant trichomes appear to be a taxa. The advent of new sequencing technologies is helping to elucidate the microbial diversity in many ecosystems, but more studies are needed to document the functional role of fungi in the microbial communities thriving in these unusual environments.     

Potent toxins in Arctic environments – Presence of saxitoxins and an unusual microcystin variant in Arctic freshwater ecosystems

Cyanobacteria are the predominant phototrophs in freshwater ecosystems of the polar regions where they commonly form extensive benthic mats. Despite their major biological role in these ecosystems, little attention has been paid to their physiology and biochemistry. An important feature of cyanobacteria from the temperate and tropical regions is the production of a large variety of toxic secondary metabolites. In Antarctica, and more recently in the Arctic, the cyanobacterial toxins microcystin and nodularin (Antarctic only) have been detected in freshwater microbial mats. To date other cyanobacterial toxins have not been reported from these locations. Five Arctic cyanobacterial communities were screened for saxitoxin, another common cyanobacterial toxin, and microcystins using immunological, spectroscopic and molecular methods. Saxitoxin was detected for the first time in cyanobacteria from the Arctic. In addition, an unusual microcystin variant was identified using liquid chromatography–mass spectrometry. Gene expression analyses confirmed the analytical findings, whereby parts of the sxt and mcy operon involved in saxitoxin and microcystin synthesis, were detected and sequenced in one and five of the Arctic cyanobacterial samples, respectively. The detection of these compounds in the cryosphere improves the understanding of the biogeography and distribution of toxic cyanobacteria globally. The sequences of sxt and mcy genes provided from this habitat for the first time may help to clarify the evolutionary origin of toxin production in cyanobacteria.     

Recent research on Arctic benthos: common notions need to be revised

Increased public awareness of the global significance of polar regions and opening of the Russian Arctic to foreign researchers have led to a pronounced intensification of benthic research in Arctic seas. The wealth of information gathered in these efforts has markedly enhanced our knowledge on the Arctic benthos. While some scientific concepts have been corroborated by the novel findings (e.g., low endemism and high faunistic affinity to northern Atlantic assemblages), other common notions need to be revised, particularly with regard to the often-cited differences between Arctic seas and the Southern Ocean. It has been demonstrated that benthos assemblages vary broadly in diversity between Arctic regions and that, hence, the idea of a consistently poor Arctic benthos—being in stark contrast to the rich Antarctic bottom fauna—is an undue overgeneralization. In terms of biogeographic diversity, both Arctic and Antarctic waters seem to be characterized by intermediate species richness. Levels of disturbance—a major ecological agent known to heavily affect benthic diversity and community structure—have been assumed to be relatively high in the Arctic but exceptionally low in the Southern Ocean. The discovery of the great role of iceberg scouring in Antarctic shelf ecosystems, which has largely been overlooked in the past, calls for a reconsideration of this notion. The novel data clearly demonstrate that there are marked differences in geographical and environmental setting, impact of fluvial run-off, pelagic production regime, strength of pelago–benthic coupling and, hence, food supply to the benthos among the various Arctic seas, impeding the large-scale generalization of local and regional findings. Field evidence points to the great significance of meso-scale features in hydrography and ice cover (marginal ice zones, polynyas, and gyres) as ‘hot spots’ of tight pelago–benthic coupling and, hence, high benthic biomass. In contrast, the importance of terrigenic organic matter discharged to the Arctic seas through fluvial run-off as an additional food source for the benthos is still under debate. Studies on the partitioning of energy flow through benthic communities strongly suggest that megafauna has to be adequately considered in overall benthic energy budgets and models of carbon cycling, particularly in Arctic shelf systems dominated by abundant echinoderm populations. Much progress has been made in the scientific exploration of the deep ice-covered Arctic Ocean. There is now evidence that it is one order of magnitude more productive than previously thought. Therefore, the significance of shelf–basin interactions, i.e., the importance of excess organic carbon exported from productive shelves to the deep ocean, is still debated and, hence, a major topic of on-going research. Another high-priority theme of current/future projects are the ecological consequences of the rapid warming in the Arctic. Higher water temperatures, increased fluvial run-off and reduced ice cover will give rise to severe ecosystem changes, propagating through all trophic levels. It is hypothesized that there would be a shift in the relative importance of marine biota in the overall carbon and energy flux, ultimately resulting in a switch from a ‘sea-ice algae–benthos’ to a ‘phytoplankton–zooplankton’ dominance.     

Psychrophilic versus psychrotolerant bacteria--occurrence and significance in polar and temperate marine habitats.

The numerical dominance and ecological role of psychrophilic bacteria in bottom sediments, sea ice, surface water and melt pools of the polar oceans were investigated using isolates, colony forming units (CFU) and metabolic activities. All sediment samples of the Southern Ocean studied showed a clear numerical dominance of cold-loving bacteria. In Arctic sediments underlying the influence of cold polar water bodies psychrophiles prevailed also but they were less dominant in sediments influenced by the warm Atlantic Water. A predominance of psychrophiles was further found in consolidated Antarctic sea ice as well as in multiyear Arctic sea ice and in melt pools on top of Arctic ice floes. A less uniform adaptation response was, however, met in polar surface waters. In the very northern part of the Fram Strait (Arctic Ocean) we found bacterial counts and activities at 1 degree C exceeding those at 22 degrees C. In surface water of the Weddell Sea (Southern Ocean) psychrophiles also dominated numerically in early autumn but the dominance declined obviously with the onset of winter-water and a decrease of chlorphyll a. Otherwise in surface water of the Southern Ocean CFUs were higher at 22 degrees C than at 1 degree C while activities were vice versa indicating at least a functional dominance of psychrophiles. Even in the temperate sediments of the German Bight true psychrophiles were present and a clear shift towards cold adapted communities in winter observed. Among the polar bacteria a more pronounced cold adaptation of Antarctic in comparison with Arctic isolates was obtained. The results and literature data indicate that stenothermic cold adapted bacteria play a significant role in the global marine environment. On the basis of the temperature response of our isolates from different habitats it is suggested to expand the definition of Morita in order to meet the cold adaptation strategies of the bacteria in the various cold habitats.     

Mycorrhizas and dark septate root endophytes in polar regions

We review the distributions and functions of mycorrhizas and dark septate root endophytes in polar regions. Arbuscular mycorrhizas (AM) are present in the Arctic and Antarctic to 82 °N and 63 °S, respectively, with fine endophyte being the dominant form of AM in roots at higher latitudes. Ecto- (ECM) and ericoid (ERM) mycorrhizas both occur in the Arctic to 79 °N, owing to the presence of species of Salix, Dryas, Vaccinium and Cassiope to this latitude. ECM and ERM are not present in Antarctic ecosystems, owing to an absence of suitable hosts. Arbutoid and orchid mycorrhizas are infrequent in the Arctic, whilst the latter are present at one location in the sub-Antarctic. Data from studies of AM, ECM and ERM colonisation along a latitudinal transect through the Arctic indicate that the frequency of plant species not colonised by mycorrhizas increases at higher latitudes, largely owing to an increase in non-mycorrhizal and a decrease in obligately mycorrhizal plant families at more northerly locations. A separate group of root- and rhizoid-associated fungi, the dark septate root endophytes (DSE), are widespread to 82 °N and 77 °S, and are apparently more frequent than mycorrhizal fungi in polar regions. The functions of DSE are largely unclear, but studies suggest beneficial effects on plant growth under defined conditions. We advocate further research into the effects of DSE on their host plants in polar regions.     

Evolution of the Arctic Calanus complex: an Arctic marine avocado?

Before man hunted the large baleen whales to near extinction by the end of the nineteenth century, Arctic ecosystems were strongly influenced by these large predators. Their main prey were zooplankton, among which the calanoid copepod species of the genus Calanus, long considered key elements of polar marine ecosystems, are particularly abundant. These herbivorous zooplankters display a range of adaptations to the highly seasonal environments of the polar oceans, most notably extensive energy reserves and seasonal migrations to deep waters where the non-feeding season is spent in diapause. Classical work in marine ecology has suggested that slow growth, long lifespan and large body size in zooplankton are specific adaptations to life in cold waters with short and unpredictable feeding seasons. Here, we challenge this understanding and, by using an analogy from the evolutionary and contemporary history of the avocado, argue that predation pressure by the now nearly extinct baleen whales was an important driving force in the evolution of life history diversity in the Arctic Calanus complex.