Phylogenetic composition of bacterioplankton assemblages from the Arctic Ocean

We analyzed the phylogenetic composition of bacterioplankton assemblages in 11 Arctic Ocean samples collected over three seasons (winter-spring 1995, summer 1996, and summer-fall 1997) by sequencing cloned fragments of 16S rRNA genes. The sequencing effort was directed by denaturing gradient gel electrophoresis (DGGE) screening of samples and the clone libraries. Sequences of 88 clones fell into seven major lineages of the domain Bacteria: alpha(36%)-, gamma(32%)-, delta(14%)-, and epsilon(1%)-Proteobacteria; Cytophaga-Flexibacter-Bacteroides spp. (9%); Verrucomicrobium spp. (6%); and green nonsulfur bacteria (2%). A total of 34% of the cloned sequences (excluding clones in the SAR11 and Roseobacter groups) had sequence similarities that were <94% compared to previously reported sequences, indicating the presence of novel sequences. DGGE fingerprints of the selected samples showed that most of the bands were common to all samples in all three seasons. However, additional bands representing sequences related to Cytophaga and Polaribacter species were found in samples collected during the summer and fall. Of the clones in a library generated from one sample collected in spring of 1995, 50% were the same and were most closely affiliated (99% similarity) with Alteromonas macleodii, while 50% of the clones in another sample were most closely affiliated (90 to 96% similarity) with Oceanospirillum sp. The majority of the cloned sequences were most closely related to uncultured, environmental sequences. Prominent among these were members of the SAR11 group. Differences between mixed-layer and halocline samples were apparent in DGGE fingerprints and clone libraries. Sequences related to alpha-Proteobacteria (dominated by SAR11) were abundant (52%) in samples from the mixed layer, while sequences related to gamma-proteobacteria were more abundant (44%) in halocline samples. Two bands corresponding to sequences related to SAR307 (common in deep water) and the high-G+C gram-positive bacteria were characteristic of the halocline samples.     

Effect of ice melting on bacterial carbon fluxes channelled by viruses and protists in the Arctic Ocean

During the last few years, extensive sea ice melting in the Arctic due to climate change has been detected, which could potentially modify the organic carbon fluxes in these waters. In this study, the effect of sea ice melting on bacterial carbon channelling by phages and protists has been evaluated in the northern Greenland Sea and Arctic Ocean. Grazing on bacteria by protists was evaluated using the FLB disappearance method. Lysis of bacteria due to viral infections was measured using the virus reduction approach. Losses of bacterial production caused by protists (PMM_BP) dominated losses caused by viruses (VMM_BP) throughout the study. Lysogenic viral production was detected in 7 out of 21 measurements and constituted from 33.9 to 100.0% of the total viral production. Significantly higher PMM_BP and lower VMM_BP were detected in waters affected by ice melting compared with unaffected waters. Consequently, significantly more bacterial carbon was channelled to the higher trophic levels in affected waters (13.05 ± 5.98 μgC l⁻¹ day⁻¹) than in unaffected waters (8.91 ± 8.33 μgC l⁻¹ day⁻¹). Viruses channelled 2.63 ± 2.45 μgC l⁻¹ day⁻¹ in affected waters and 4.27 ± 5.54 μgC l⁻¹ day⁻¹ in unaffected waters. We conclude that sea ice melting in the Arctic could modify the carbon flow through the microbial food web. This process may be especially important in the case of massive sea ice melting due to climate change.     

Metabolic diversity of heterotrophic bacterioplankton over winter and spring in the coastal Arctic Ocean

Metabolic diversity of heterotrophic bacterioplankton was tracked from early winter through spring with Biolog Ecoplates under the seasonally ice covered arctic shelf in the Canadian Arctic (Franklin Bay, Beaufort Sea). Samples were taken every 6 days from December 2003 to May 2004 at the surface, the halocline where a temperature inversion occurs, and at 200 m, close to the bottom. Despite the low nutrient levels and low chlorophyll a , suggesting oligotrophy in the winter surface waters, the number of substrates used (NSU) was greater than in spring, when chlorophyll a concentrations increased. Denaturing gradient gel electrophorisis analysis also indicated that the winter and spring bacterial communities were phylogenetically distinct, with several new bands appearing in spring. In spring, the bacterial community would have access to the freshly produced organic carbon from the early phytoplankton bloom and the growth of rapidly growing specialist phenotypes would be favoured. In contrast, in winter bacterioplankton consumed more complex organic matter originated during the previous year's phytoplankton production. At the other depths we tested the NSU was similar to that for the winter surface, with no seasonal pattern. Instead, bacterioplankton metabolism seemed to be influenced by resuspension, advection, and sedimentation events that contributed organic matter that enhanced bacterial metabolism.     

Gelatinous zooplankton of the Arctic Ocean: in situ observations under the ice

The gelatinous zooplankton of the Canada Basin were observed with a deep-diving remotely operated vehicle (ROV) during August–September, 2002. Taxa observed fell into four main groups: cnidarians, ctenophores, chaetognaths, and pelagic tunicates. We provide detailed data on the vertical distributions of many taxa from three sites which span the Canada Basin. The most common gelatinous organisms in the surface waters were the ctenophores Mertensia ovum and Bolinopsis infundibulum. These two species were found in very large numbers in the near-surface mixed layer. In the mesopelagic zone, below the transition from the Pacific water layer and the Atlantic water layer, the most common species was Sminthea arctica. Surprising numbers of the scyphomedusa Atolla tenella were found in the deep waters of the basin, along with an undescribed species of narcomedusae. The vertical distributions of the gelatinous zooplankton observed with the ROV show several trends related to the physical properties of the water and geographic location within the basin.     

Zooplankton boom and ice amphipod bust below melting sea ice in the Amundsen Gulf, Arctic Canada

Early summer in the Arctic with extensive ice melt and break-up represents a dramatic change for sympagic–pelagic fauna below seasonal sea ice. As part of the International Polar Year-Circumpolar Flaw Lead system study (IPY-CFL), this investigation quantified zooplankton in the meltwater layer below landfast ice and remaining ice fauna below melting ice during June (2008) in Franklin Bay and Darnley Bay, Amundsen Gulf, Canada. The ice was in a state of advanced melt, with fully developed melt ponds. Intense melting resulted in a 0.3- to 0.5-m-thick meltwater layer below the ice, with a strong halocline to the Arctic water below. Zooplankton under the ice, in and below the meltwater layer, was sampled by SCUBA divers. Dense concentrations (max. 1,400 ind. m⁻³) of Calanus glacialis were associated with the meltwater layer, with dominant copepodid stages CIV and CV and high abundance of nauplii. Less abundant species included Pseudocalanus spp., Oithona similis and C. hyperboreus. The copepods were likely feeding on phytoplankton (0.5–2.3 mg Chl-a m⁻³) in the meltwater layer. Ice amphipods were present at low abundance (<10 ind. m⁻²) and wet biomass (<0.2 g m⁻²). Onisimus glacialis and Apherusa glacialis made up 64 and 51% of the total ice faunal abundance in Darnley Bay and Franklin Bay, respectively. During early summer, the autochthonous ice fauna becomes gradually replaced by allochthonous zooplankton, with an abundance boom near the meltwater layer. The ice amphipod bust occurs during late stages of melting and break-up, when their sympagic habitat is diminished then lost.     

Sea ice phenology and timing of primary production pulses in the Arctic Ocean

Arctic organisms are adapted to the strong seasonality of environmental forcing. A small timing mismatch between biological processes and the environment could potentially have significant consequences for the entire food web. Climate warming causes shrinking ice coverage and earlier ice retreat in the Arctic, which is likely to change the timing of primary production. In this study, we test predictions on the interactions among sea ice phenology and production timing of ice algae and pelagic phytoplankton. We do so using the following (1) a synthesis of available satellite observation data; and (2) the application of a coupled ice‐ocean ecosystem model. The data and model results suggest that, over a large portion of the Arctic marginal seas, the timing variability in ice retreat at a specific location has a strong impact on the timing variability in pelagic phytoplankton peaks, but weak or no impact on the timing of ice‐algae peaks in those regions. The model predicts latitudinal and regional differences in the timing of ice algae biomass peak (varying from April to May) and the time lags between ice algae and pelagic phytoplankton peaks (varying from 45 to 90 days). The correlation between the time lag and ice retreat is significant in areas where ice retreat has no significant impact on ice‐algae peak timing, suggesting that changes in pelagic phytoplankton peak timing control the variability in time lags. Phenological variability in primary production is likely to have consequences for higher trophic levels, particularly for the zooplankton grazers, whose main food source is composed of the dually pulsed algae production of the Arctic.     

Arctic Ocean microbial community structure before and after the 2007 record sea ice minimum

Increasing global temperatures are having a profound impact in the Arctic, including the dramatic loss of multiyear sea ice in 2007 that has continued to the present. The majority of life in the Arctic is microbial and the consequences of climate-mediated changes on microbial marine food webs, which are responsible for biogeochemical cycling and support higher trophic levels, are unknown. We examined microbial communities over time by using high-throughput sequencing of microbial DNA collected between 2003 and 2010 from the subsurface chlorophyll maximum (SCM) layer of the Beaufort Sea (Canadian Arctic). We found that overall this layer has freshened and concentrations of nitrate, the limiting nutrient for photosynthetic production in Arctic seas, have decreased. We compared microbial communities from before and after the record September 2007 sea ice minimum and detected significant differences in communities from all three domains of life. In particular, there were significant changes in species composition of Eukarya, with ciliates becoming more common and heterotrophic marine stramenopiles (MASTs) accounting for a smaller proportion of sequences retrieved after 2007. Within the Archaea, Marine Group I Thaumarchaeota, which earlier represented up to 60% of the Archaea sequences in this layer, have declined to <10%. Bacterial communities overall were less diverse after 2007, with a significant decrease of the Bacteroidetes. These significant shifts suggest that the microbial food webs are sensitive to physical oceanographic changes such as those occurring in the Canadian Arctic over the past decade.     

rRNA and rDNA based assessment of sea ice protist biodiversity from the central Arctic Ocean

Sea ice is a large and diverse ecosystem contributing significantly to primary production in ice-covered regions. In the Arctic Ocean, sea ice consists of mixed multi-year ice (MYI), often several metres thick, and thinner first-year ice (FYI). Current global warming is most severe in Arctic regions; as a consequence, summer sea ice cover is decreasing and MYI is disappearing at an alarming rate. Despite its apparent hostility, sea ice is inhabited by a diverse microbial community of bacteria and protists, many of which are photosynthetic. Here we present an assessment of eukaryotic biodiversity in MYI and FYI from the central Arctic Ocean using high-throughput 454 sequencing of 18S rRNA and rDNA amplicons. We compared the rDNA-based ‘total’ biodiversity with the ‘active’ biodiversity from rRNA amplicons and found differences between them including an over-representation of Ciliophora, Bicosoecida and Bacillariophyceae operational taxonomic units (OTUs) in the active part of the community. Differences between the two libraries are more pronounced at the lower taxonomic level: certain genera, such as Melosira, are more abundant in the rRNA library, indicating activity of these genera. Furthermore, we found that one FYI station showed a higher activity of potential grazers which was probably due to the advanced stage of melt evident by higher ice temperatures and highly porous ice compared with the other stations.     

High viral infection rates in Antarctic and Arctic bacterioplankton

The frequency of visibly phage-infected bacterial cells (FVIB) and the average number of phages per cell [i.e. burst size (BS)] were determined in Antarctic and Arctic ultra-oligotrophic freshwater environments. Water samples were collected from two Antarctic freshwater lakes and cryoconite holes from a glacier in the Arctic. Data from this bipolar study show the highest FVIB (average 26.1%, range 5.1% to 66.7%) and the lowest BS (average 4, range 2-15) ever reported in the literature. The bacterial density is low in these ultra-oligotrophic freshwater environments but a large proportion of the bacteria are visibly infected. Our results suggest that a constant virioplankton population can be maintained in these extreme environments even though host density is low and often slow growing.     

Impacts of Ocean Acidification on Sediment Processes in Shallow Waters of the Arctic Ocean

Despite the important roles of shallow-water sediments in global biogeochemical cycling, the effects of ocean acidification on sedimentary processes have received relatively little attention. As high-latitude cold waters can absorb more CO₂ and usually have a lower buffering capacity than warmer waters, acidification rates in these areas are faster than those in sub-tropical regions. The present study investigates the effects of ocean acidification on sediment composition, processes and sediment-water fluxes in an Arctic coastal system. Undisturbed sediment cores, exempt of large dwelling organisms, were collected, incubated for a period of 14 days, and subject to a gradient of pCO₂ covering the range of values projected for the end of the century. On five occasions during the experimental period, the sediment cores were isolated for flux measurements (oxygen, alkalinity, dissolved inorganic carbon, ammonium, nitrate, nitrite, phosphate and silicate). At the end of the experimental period, denitrification rates were measured and sediment samples were taken at several depth intervals for solid-phase analyses. Most of the parameters and processes (i.e. mineralization, denitrification) investigated showed no relationship with the overlying seawater pH, suggesting that ocean acidification will have limited impacts on the microbial activity and associated sediment-water fluxes on Arctic shelves, in the absence of active bio-irrigating organisms. Only following a pH decrease of 1 pH unit, not foreseen in the coming 300 years, significant enhancements of calcium carbonate dissolution and anammox rates were observed. Longer-term experiments on different sediment types are still required to confirm the limited impact of ocean acidification on shallow Arctic sediment processes as observed in this study.     

北极浮冰生态学研究进展

近年来随着北极地区的开放和全球变化对北极地区生态环境和海冰现存量的影响日益显现,北极浮冰生态学研究得到了广泛的重视和实质性的进展.最新研究结果显示,浮冰本身包含了一个复杂的生物群落,高纬度浮冰生物群落的初级产量远高于原先的估算,浮冰生物群落在北极海洋生态系统中的作用被进一步确认.但由于对浮冰生物群落的研究受后勤保障条件的制约,目前尚有大量科学问题有待今后进一步深入研究,预期我国科学家将在其中做出贡献.     

Virus impact on heterotrophic bacterioplankton of water reservoirs

The quantitative distribution of viruses and their impact on heterotrophic bacterioplankton were studied in mesotrophic and eutrophic reservoirs of the Volga and Volga-Baltic waterway. The abundance of planktonic virus particles ranged from 9.4 × 10⁶ to 120 × 10⁶ ml⁻¹ and was from 2.5 to 9 times greater than the bacterial numbers. Production of virioplankton varied from 2.1 × 10⁶ to 132 × 10⁶ particles (ml day)⁻¹ and the population turnover time values were between 0.3 and 11.6 days. The maximum values of numbers and production of virio- and bacterioplankton were observed in the eutrophic Ivan’kovo reservoir. Distribution of the viruses in the Volga reservoirs depended to a significant degree on the number and activity of heterotrophic bacterioplankton. The infected bacteria accounted for 5.5–33.5% of the total bacterial abundance. Phages were an important factor of bacterial mortality. During July to September virus-induced bacterial mortality varied between 6.1 and 40.6% (20.2% on average) of daily bacterioplankton production.     

Relevance of bacterioplankton abundance and production in the oligotrophic equatorial Indian Ocean

Bacterioplankton abundance and production, chlorophyll a (Chl a) concentrations and primary production (PP) were measured from the equatorial Indian Ocean (EIO) during northeast (NEM), southwest (SWM) and spring intermonsoon (SpIM) seasons from 1°N to 5°S along 83°E. The average bacterial abundance was 0.52 ± 0.29, 0.62 ± 0.33 and 0.46 ± 0.19 (× 10⁸ cells l⁻¹), respectively during NEM, SWM and SpIM in the top 100 m. In the deep waters (200 m and below), the bacterial counts averaged ∼0.35 ± 0.14 × 10⁸ cells l⁻¹ in SWM and 0.39 ± 0.16 × 10⁸ cells l⁻¹ in SpIM. The 0–120 m column integrated bacterial production (BP) ranged from 19 to 115 and from 10 to 51 mg C m⁻² d⁻¹ during NEM and SWM, respectively. Compared with many open ocean locations, bacterial abundance and production in this region are lower. The bacterial carbon production, however, is notably higher than that of phytoplankton PP (BP:PP ratio 102% in SWM and 188% in NEM). With perpetually low PP (NEM: 20, SWM: 18 and SpIM: 12 mg C m⁻² d⁻¹) and Chl a concentration (NEM: 16.5, SWM: 15.0 and SpIM: 20.9 mg m⁻²), the observed bacterial abundance and production are pivotal in the trophodynamics of the EIO. Efficient assimilation and mineralization of available organics by bacteria in the euphotic zone might serve a dual role in the ultra-oligotrophic regions including EIO. Thus, bacteria probably sustain microheterotrophs (micro- and meso-zooplankton) through microbial loop. Further, rapid mineralization by bacteria will make essential nutrients available to autotrophs.     

The role of sea ice for vascular plant dispersal in the Arctic

Sea ice has been suggested to be an important factor for dispersal of vascular plants in the Arctic. To assess its role for postglacial colonization in the North Atlantic region, we compiled data on the first Late Glacial to Holocene occurrence of vascular plant species in East Greenland, Iceland, the Faroe Islands and Svalbard. For each record, we reconstructed likely past dispersal events using data on species distributions and genetics. We compared these data to sea-ice reconstructions to evaluate the potential role of sea ice in these past colonization events and finally evaluated these results using a compilation of driftwood records as an independent source of evidence that sea ice can disperse biological material. Our results show that sea ice was, in general, more prevalent along the most likely dispersal routes at times of assumed first colonization than along other possible routes. Also, driftwood is frequently dispersed in regions that have sea ice today. Thus, sea ice may act as an important dispersal agent. Melting sea ice may hamper future dispersal of Arctic plants and thereby cause more genetic differentiation. It may also limit the northwards expansion of competing boreal species, and hence favour the persistence of Arctic species.     

Notes on the biology of sea ice in the Arctic and Antarctic

The sea ice which covers large areas of the polar regions plays a major role in the marine ecosystem of both the Arctic and Southern Oceans. Not only do warmblooded animals depend on sea ice as a platform, but the sympagic organisms living internally within the sea ice or at the interfaces ice/snow and ice/water provide a substantial part of the total primary production of the ice covered regions. In addition sea ice organisms are an important food source for a variety of pelagic animals and may initiate phytoplankton spring blooms after ice melt by seeding effects.Sea ice organisms often are enriched by some orders of magnitude if the same volume of melted ice is compared to that of the underlying water column. Three hypotheses try to explain this discrepancy and are discussed. Investigations on the nutrient chemistry within the sea ice system and in-situ observations still are rare. Intense growth of sympagic organisms can result in nutrient deficiencies, at least in selected habitats. Advances in endoscopie methods may lead to a better understanding of the life within the sea ice.Paper presented at the Symposium on Polar regions: the challenge for biological and ecological research organised by the Swiss Committee for Polar Research, Basel on 2 October 1992     

北冰洋海域微食物环研究进展

海洋微食物环在海洋生态系统中起着重要作用.北冰洋因常年为海冰所覆盖,对微食物环的研究较为有限.现有研究表明,微食物环在北冰洋生态系统中的作用与海域和季节相关.近年来环境的快速变化、特别是夏季海冰覆盖面积的迅速减少,会对微食物环的结构和功能产生重大影响,已有研究显示其生态作用有望进一步提高.综合近年来已有的研究成果,对北冰洋微食物环的主要类群:原核生物、真核浮游植物、原生动物和浮游病毒等的基本生态特征进行了概述,讨论了各类群间的相互关系,并对未来的研究重点进行了展望.     

Population dynamics of bacteria in Arctic sea ice

The dynamics of bacterial populations in annual sea ice were measured throughout the vernal bloom of ice algae near Resolute in the Canadian Arctic. The maximum concentration of bacteria was 6.0·10¹¹ cells·m^–2 (about 2.0·10¹⁰ cells·l^–1) and average cell volume was 0.473 m³ in the lower 4 cm of the ice sheet. On average, 37% of the bacteria were epiphytic and were most commonly attached (70%) to the dominant alga,Nitzschia frigida (58% of total algal numbers). Bacterial population dynamics appeared exponential, and specific growth rates were higher in the early season (0.058 day^–1), when algal biomass was increasing, than in the later season (0.0247 day^–1), when algal biomass was declining. The proportion of epiphytes and the average number of epiphytes per alga increased significantly (P<0.05) through the course of the algal bloom. The net production of bacteria was 67.1 mgC·m^–2 throughout the algal bloom period, of which 45.5 mgC·m^–2 occurred during the phase of declining algal biomass. Net algal production was 1942 mgC·m^–2. Sea ice bacteria (both arctic and antarctic) are more abundant than expected on the basis of relationships between bacterioplankton and chlorophyll concentrations in temperate waters, but ice bacteria biomass and net production are nonetheless small compared with the ice algal blooms that presumably support them.     

Seasonal changes of Antarctic marine bacterioplankton and sea ice bacterial assemblages

The distributions of bacterial populations in sea ice and underlying seawater were investigated on the continental shelf of the “Terre Adélie” area. A reference station was sampled weekly from January 1991 to January 1992. In winter, the survey included a minimum of six sampling layers: surface and bottom ice, brine, seawater from the interface, and at 0.5 and 2 m depth. In seawater, the total bacterial abundance ranged from 0.5 × 10⁵ cells ml⁻¹ in July to 6.0 × 10⁵ cells ml⁻¹ after ice break. Values reaching 2.5 × 10⁶ cells ml⁻¹ were recorded in the overlying ice cover. Mean cell volumes were twice as high in brine as in seawater. The saprophytic bacterial abundance ranged from 5.0 × 10⁴ CFU (colony-forming units) ml⁻¹ in some winter interface samples to less than 1.0 × 10³ CFU ml⁻¹ in most of the summer seawater samples. In sea ice a clear decreasing gradient for most of the studied bacterial parameters from the surface layers towards the bottom layer was found. The ice cover had a discernible impact on underlying seawater, but its influence was restricted to a limited interface layer.