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Due to climate change, increased microbial activity in high-latitude soils may lead to higher greenhouse gas (GHG) emissions. However, microbial GHG production and consumption mechanisms in tundra soils are not thoroughly understood. To investigate how the diversity and functional potential of bacterial and archaeal communities vary across vegetation types and soil layers, we analyzed 116 soil metatranscriptomes from 73 sites in the Finnish sub-Arctic. Meadow soils were characterized by higher pH and lower soil organic matter (SOM) and carbon/nitrogen ratio. By contrast, dwarf shrub-dominated ecosystems had higher SOM and lower pH. Although Actinobacteria, Acidobacteria, Alphaproteobacteria and Planctomycetes were dominant in all communities, there were significant differences at the genus level between vegetation types; plant polymer-degrading groups were more active in shrub-dominated soils than in meadows. Given that climate-change scenarios predict the expansion of shrubs at high latitudes, our results indicate that tundra soil microbial communities harbor potential decomposers of increased plant litter, which may affect the rate of carbon turnover in tundra soils. Additionally, transcripts of methanotrophs were detected in the mineral layer of all soils, which may moderate methane fluxes. This study provides new insights into possible shifts in tundra microbial diversity and activity due to climate change. Active microbial communities were significantly different in the organic and mineral soil layers and the communities differed significantly between four different vegetation types both in the organic and mineral layers.

Poly-3-hydroxyalkanoic acids (PHAs) are bacterial storage polymers commonly used in bioplastic production. Halophilic bacteria are industrially interesting organisms, as their salinity tolerance and psychrophilic nature lowers sterility requirements and subsequent production costs. We investigated PHA synthesis in two bacterial strains, Halomonas sp. 363 and Paracoccus sp. 392, isolated from Southern Ocean sea ice and elucidated the related PHA biopolymer accumulation and composition with various approaches, such as transcriptomics, microscopy, and chromatography. We show that both bacterial strains produce PHAs at 4 degrees C when the availability of nitrogen and/or oxygen limited growth. The genome of Halomonas sp. 363 carries three phaC synthase genes and transcribes genes along three PHA pathways (I to III), whereas Paracoccus sp. 392 carries only one phaC gene and transcribes genes along one pathway (I). Thus, Halomonas sp. 363 has a versatile repertoire of phaC genes and pathways enabling production of both short- and medium-chain-length PHA products. IMPORTANCE Plastic pollution is one of the most topical threats to the health of the oceans and seas. One recognized way to alleviate the problem is to use degradable bioplastic materials in high-risk applications. PHA is a promising bioplastic material as it is nontoxic and fully produced and degraded by bacteria. Sea ice is an interesting environment for prospecting novel PHA-producing organisms, since traits advantageous to lower production costs, such as tolerance for high salinities and low temperatures, are common. We show that two sea-ice bacteria, Halomonas sp. 363 and Paracoccus sp. 392, are able to produce various types of PHA from inexpensive carbon sources. Halomonas sp. 363 is an especially interesting PHA-producing organism, since it has three different synthesis pathways to produce both short- and medium-chain-length PHAs. peerReviewed

Microbial consumption of phytoplankton-derived organic carbon in the pelagic food web is an important component of the global C cycle. We studied C cycling in two phytoplankton-bacteria systems (non-axenic cultures of a dinoflagellate Apocalathium malmogiense and a cryptophyte Rhodomonas marina) in two experiments. In the first experiment we grew phytoplankton and bacteria in nutrient replete conditions and followed C processing at early exponential growth phase and at two later phases. Primary production and total community respiration were up to 4 and 7 times higher, respectively, in the A. malmogiense treatments. Based on the optical signals, accumulating dissolved organic C (DOC) was degraded more in the R. marina treatments and the rate of bacterial production to primary production was higher. Thus, the flow of C from phytoplankton to bacteria was relatively higher in R. marina treatments than in A. malmogiense treatments which was further supported by faster 14C transfer from phytoplankton to bacterial biomass. In the second experiment we investigated consumption of the phytoplankton-derived DOC by bacteria. DOC consumption and transformation, bacterial production and bacterial respiration were all higher in R. marina treatments. In both experiments A. malmogiense supported a bacterial community predominated by bacteria specialized in the utilization of less labile DOC (class Bacteroidia) whereas R. marina supported a community predominated by copiotrophic Alpha- and Gammaproteobacteria. Our findings suggest that large dinoflagellates cycle relatively more C between phytoplankton biomass and the inorganic C pool whereas small cryptophytes direct relatively more C to the microbial loop.

Increased microbial activity in high-latitude soils due to climate change might lead to higher greenhouse gas (GHG) emissions. However, mechanisms of microbial GHG production and consumption in tundra soils are not thoroughly understood. We analyzed 116 soil metatranscriptomes from 73 sites in the Finnish sub-Arctic to investigate how the diversity and functional potential of bacterial and archaeal communities vary across vegetation types and soil layers. Soils differed in physicochemical conditions, with meadow soils being characterized by higher pH and low soil organic matter (SOM) and carbon/nitrogen ratio whereas dwarf shrub-dominated ecosystems with high SOM and low pH. Actinobacteria, Acidobacteria, Alphaproteobacteria, and Planctomycetes predominated all communities but there were significant differences on genus level between vegetation types, as plant polymer degrading groups were more active in shrub-dominated soils compared to meadows. Given that climate change scenarios predict expansion in dwarf shrubs at high latitudes, our results indicate that the rate of carbon turnover in tundra soils may increase in the future. Additionally, transcripts of methanotrophs were detected in the mineral layer of all soils, potentially moderating methane fluxes from deeper layers. In all, this study provides new insights into possible shifts in tundra microbial diversity and activity with climate change.

Although studies of biogeochemical processes in polar sea ice have been increasing, similar research on relatively warm low-salinity sea ice remains sparse. In this study, we investigated biogeochemical properties of the landfast sea ice cover in the brackish Bothnian Bay (Northern Baltic Sea) and the possible role of this sea ice in mediating the exchange of greenhouse gases, including carbon dioxide (CO2), methane (CH4), and nitrous oxide (N2O) across the water column–sea ice–atmosphere interface. Observations of total alkalinity and dissolved inorganic carbon in both landfast sea ice and the water column suggest that the carbonate system is mainly driven by salinity. While high CH4 and N2O concentrations were observed in both the water column (up to 14.3 and 17.5 nmol L–1, respectively) and the sea ice (up to 143.6 and 22.4 nmol L–1, respectively),these gases appear to be enriched in sea ice compared to the water column.This enrichment may be attributable to the sea ice formation process, which concentrates impurities within brine. As sea ice temperature and brine volume decrease, gas solubility decreases as well, promoting the formation of bubbles. Gas bubbles originating from underlying sediments may also be incorporated within the ice cover and contribute to the enrichment in sea ice.The fate of these greenhouse gases within the ice merits further research, as storage in this low-salinity seasonal sea ice is temporary.

Publisher Copyright: © 2021 The Author(s). A rigorous synthesis of the sea-ice ecosystem and linked ecosystem services highlights that the sea-ice ecosystem supports all 4 ecosystem service categories, that sea-ice ecosystems meet the criteria for ecologically or biologically significant marine areas, that global emissions driving climate change are directly linked to the demise of sea-ice ecosystems and its ecosystem services, and that the sea-ice ecosystem deserves specific attention in the evaluation of marine protected area planning. The synthesis outlines (1) supporting services, provided in form of habitat, including feeding grounds and nurseries for microbes, meiofauna, fish, birds and mammals (particularly the key species Arctic cod, Boreogadus saida, and Antarctic krill, Euphausia superba, which are tightly linked to the sea-ice ecosystem and transfer carbon from sea-ice primary producers to higher trophic level fish, mammal species and humans); (2) provisioning services through harvesting and medicinal and genetic resources; (3) cultural services through Indigenous and local knowledge systems, cultural identity and spirituality, and via cultural activities, tourism and research; (4) (climate) regulating services through light regulation, the production of biogenic aerosols, halogen oxidation and the release or uptake of greenhouse gases, for example, carbon dioxide. The ongoing changes in the polar regions have strong impacts on sea-ice ecosystems and associated ecosystem services. While the response of sea-ice–associated primary production to environmental change is regionally variable, the effect on ice-associated mammals and birds is predominantly negative, subsequently impacting human harvesting and cultural services in both polar regions. Conservation can help protect some species and functions. However, the key mitigation measure that can slow the transition to a strictly seasonal ice cover in the Arctic Ocean, reduce the overall loss of sea-ice habitats from the ocean, and thus preserve the unique ecosystem services provided by sea ice and their contributions to human well-being is a reduction in carbon emissions.

Background In contrast to earlier assumptions, there is now mounting evidence for the role of tundra soils as important sources of the greenhouse gas nitrous oxide (N2O). However, the microorganisms involved in the cycling of N2O in this system remain largely uncharacterized. Since tundra soils are variable sources and sinks of N2O, we aimed at investigating differences in community structure across different soil ecosystems in the tundra. Results We analysed 1.4 Tb of metagenomic data from soils in northern Finland covering a range of ecosystems from dry upland soils to water-logged fens and obtained 796 manually binned and curated metagenome-assembled genomes (MAGs). We then searched for MAGs harbouring genes involved in denitrification, an important process driving N2O emissions. Communities of potential denitrifiers were dominated by microorganisms with truncated denitrification pathways (i.e., lacking one or more denitrification genes) and differed across soil ecosystems. Upland soils showed a strong N2O sink potential and were dominated by members of the Alphaproteobacteria such as Bradyrhizobium and Reyranella. Fens, which had in general net-zero N2O fluxes, had a high abundance of poorly characterized taxa affiliated with the Chloroflexota lineage Ellin6529 and the Acidobacteriota subdivision Gp23. Conclusions By coupling an in-depth characterization of microbial communities with in situ measurements of N2O fluxes, our results suggest that the observed spatial patterns of N2O fluxes in the tundra are related to differences in the composition of denitrifier communities.

In recent decades, the phytoplankton community in parts of the Baltic Sea has shifted from diatom dominance to co-occurrence of diatoms and dinoflagellates during the spring bloom. We investigated whether this shift affects bacterial production (BP), abundance and community composition (BCC). Two mesocosm experiments were carried out with water from the SW coast of Finland during the winters of 2012 and 2013. The water was collected before the onset of the spring bloom. Natural seawater was used as a control, and various inocula of diatom and dino flagellate cultures were used as treatments. After the phytoplankton bloom development, BP (thymidine: BPT; leucine: BPL) was significantly higher in the diatom treatments than in the controls and dinoflagellate treatments (BPT and BPL in 2012 and BPL in 2013). In 2013, the BCC was significantly different between the diatom and dinoflagellate treatments and there was a temporal shift in both experiments. Alphaproteobacteria predominated in all treatments at the beginning of the experiments and shifted to flavobacterial (2012) and betaproteobacterial predominance (2013) during the chlorophyll a peak. Towards the end of the experiment, Actinobacteria and Betaproteo bacteria predominated in the diatom treatment in 2012, whereas in 2013 Flavobacteriia (all treatments) predominated together with Gammaproteobacteria and Cytophagia (diatom treatments). The results demonstrated that bacterial physiology and community structure are affected by relatively small changes in the phytoplankton community. Thus, the ongoing changes in the phytoplankton community resulting from co-occurrence of diatoms and dinoflagellates may decrease pelagic remineralization of carbon and reduce organic matter fluxes through the microbial loop.

Horizontal and vertical variability of first-year drift-ice bacterial communities was investigated along a North-South transect in the Fram Strait during the winter/spring transition. Two different developmental stages were captured along the transect based on the prevailing environmental conditions and the differences in bacterial community composition. The differences in the bacterial communities were likely driven by the changes in sea-ice algal biomass (2.6-5.6 fold differences in chl-a concentrations). Copiotrophic genera common in late spring/summer sea ice, such as Polaribacter, Octadecabacter and Glaciecola, dominated the bacterial communities, supporting the conclusion that the increase in the sea-ice algal biomass was possibly reflected in the sea-ice bacterial communities. Of the dominating bacterial genera, Polaribacter seemed to benefit the most from the increase in algal biomass, since they covered approximately 39% of the total community at the southernmost stations with higher (>6 μg l(-1) ) chl-a concentrations and only 9% at the northernmost station with lower chl-a concentrations (

Viruses are recognized as important actors in ocean ecology and biogeochemical cycles, but many details are not yet understood. We participated in a winter expedition to the Weddell Sea, Antarctica, to isolate viruses and to measure virus-like particle abundance (flow cytometry) in sea ice. We isolated 59 bacterial strains and the first four Antarctic sea-ice viruses known (PANV1, PANV2, OANV1 and OANV2), which grow in bacterial hosts belonging to the typical sea-ice genera Paraglaciecola and Octadecabacter. The viruses were specific for bacteria at the strain level, although OANV1 was able to infect strains from two different classes. Both PANV1 and PANV2 infected 11/15 isolated Paraglaciecola strains that had almost identical 16S rRNA gene sequences, but the plating efficiencies differed among the strains, whereas OANV1 infected 3/7 Octadecabacter and 1/15 Paraglaciecola strains and OANV2 1/7 Octadecabacter strains. All the phages were cold-active and able to infect their original host at 0 degrees C and 4 degrees C, but not at higher temperatures. The results showed that virus-host interactions can be very complex and that the viral community can also be dynamic in the winter-sea ice.

1. The formation of sea ice impacts directly on the physical dynamics of water masses (e.g. wind stress at the sea surface) and air-sea exchange processes (e.g. vertical heat fluxes). 2. The annual cycle of formation, consolidation and melting of sea ice has a major influence on the ecology of both the benthic and pelagic components of the Baltic Sea ecosystem. 3. There is considerable inter-annual variation in the extent of sea ice in the Baltic Sea and thus in the size of the habitat for sympagic (ice-associated) microbial and metazoan communities as well as for larger organisms living on the ice, notably the ringed seal. 4. There is a pronounced gradient in ice characteristics, from more saline ice in the south of the Baltic Sea to freshwater ice in the north. The former is more porous and supports more ice-associated biology than the latter. 5. The Baltic sympagic communities consist mainly of prokaryotic and eukaryotic microbes (bacteria, diatoms, dinoflagellates, flagellates), ciliates and rotifers. These communities are recruited from the plankton when the ice forms, followed by an ice-adapted successional pattern with an expansion of substrate-bound pennate diatoms, which does not occur in the seawater beneath the ice. 6. The sea-ice food webs inside the ice are truncated compared to the open-water food webs because organisms larger than the upper size limit of the brine channels are lacking in the internal sympagic communities. 7. Global climate change decreases the extension and thickness of the sea ice as well as the length of the ice season, and therefore the seasonal effects that sea ice has on the Baltic Sea winter-spring ecosystem dynamics.

Polyhydroxyalkanoates (PHAs) are macromolecules produced by bacteria as means for storing carbon and energy in intracellular granules. PHAs have physical properties similar to those of plastics and have become of interest to industry as materials for environmentally friendly bioplastic production. There is an ongoing search for new PHA-producing bacterial strains and PHA-synthesizing enzymes tolerating extreme conditions to find ways of producing PHAs at cold temperatures and high solute concentrations. Moreover, the study of PHA producers in the sea-ice biome can aid in understanding the microbial ecology of carbon cycling in ice-associated ecosystems. In this study, PHA producers and PHA synthase genes were examined under the extreme environmental conditions of sea ice and cold seawater to find evidence of PHA production in an environment requiring adaptation to high salinity and cold temperatures. Sea ice and cold estuarine water samples were collected from the northern Baltic Sea and evidence of PHA production was gathered, using microscopy with Nile Blue A staining of PHA-granules and PCR assays detecting PHA-synthesis genes. The PHA granules and PHA synthases were found at all sampling locations, in both sea ice and water, and throughout the sampling period spanning over 10 years. Our study shows, for the first time, that PHA synthesis occurs in Baltic Sea cold-adapted bacteria in their natural environment, which makes the Baltic Sea and its cold environments an interesting choice in the quest for PHA-synthesizing bacteria and synthesis genes.

Antarctic sea-ice bacterial community composition and dynamics in various developmental stages were investigated during the austral winter in 2013. Thick snow cover likely insulated the ice, leading to high (

In search for sea ice bacteria and their phages from the Baltic Sea ice, two ice samples were collected from land-fast ice in a south-west Finland coastal site in February and March 2011. Bacteria were isolated from the melted sea ice samples and phages were screened from the same samples for 43 purified isolates. Plaque-producing phages were found for 15 bacterial isolates at 3 °C. Ten phage isolates were successfully plaque purified and eight of them were chosen for particle purification to analyze their morphology and structural proteins. Phage 1/32 infecting an isolate affiliated to phylum Bacteroidetes (Flavobacterium sp.) is a siphovirus and six phages infecting isolates affiliated to γ-Proteobacteria (Shewanella sp.) hosts were myoviruses. Cross titrations between the hosts showed that all studied phages are host specific. Phage solutions, host growth and phage infection were tested in different temperatures revealing phage temperature tolerance up to 45 °C, whereas phage infection was in most of the cases retarded above 15 °C. This study is the first to report isolation and cultivation of ice bacteria and cold-active phages from the Baltic Sea ice.

The dynamics of dissolved organic matter (DOM) and the succession of bacterial community composition (BCC) were investigated during bloom of filamentous cyanobacteria in a mesocosm experiment conducted in the western Gulf of Finland, the Baltic Sea. The effects of labile dissolved organic carbon (glucose), inorganic nutrients (N and P) and large zooplankton (> 100 µm) on the DOM pool, bacterial production and the composition of bacterial communities were analysed over a period of ten days. In addition, the bioavailability of dissolved organic carbon (DOC) and its turnover by heterotrophic bacteria (biomass and respiration) were investigated in three one-week bacterial bioassays. Heterotrophic bacteria rapidly utilised about 25-55 % of the DOC released from the plankton community, thus assuming it to be highly labile DOC. More than half of the accumulating net DOC pool was degraded over seven days, thus assuming it to be labile. In average, labile autochthonous DOC was degraded with bacterial growth efficiency of 25%. A distinct succession of bacterial communities accompanied the supply of autochthonous DOM, with the most prominent responses occurring in a few single phylotypes of the Delta- and Gammaproteobacterial classes. About 40% of the variation in the relative shares of dominant bacterial classes could be explained by changes in the functional groups of autotrophs. Inorganic nutrient treatment proved beneficial to Deltaproteobacteria and increased bacterial production over that of other mesocosms.

Horizontal and vertical variability of first-year drift-ice bacterial communities was investigated along a North-South transect in the Fram Strait during the winter/spring transition. Two different developmental stages were captured along the transect based on the prevailing environmental conditions and the differences in bacterial community composition. The differences in the bacterial communities were likely driven by the changes in sea-ice algal biomass (2.6-5.6 fold differences in chl-a concentrations). Copiotrophic genera common in late spring/summer sea ice, such as Polaribacter, Octadecabacter and Glaciecola, dominated the bacterial communities, supporting the conclusion that the increase in the sea-ice algal biomass was possibly reflected in the sea-ice bacterial communities. Of the dominating bacterial genera, Polaribacter seemed to benefit the most from the increase in algal biomass, since they covered approximately 39% of the total community at the southernmost stations with higher (6 μg l(-1) ) chl-a concentrations and only 9% at the northernmost station with lower chl-a concentrations (6 μg l(-1) ). The sea-ice bacterial communities also varied between the ice horizons at all three stations and thus we recommend that for future studies multiple ice horizons be sampled to cover the variability in sea-ice bacterial communities in spring.

Drift ice, open water and under-ice water bacterial communities covering several developmental stages from open water to thick ice were studied in the northern Baltic Sea. The bacterial communities were assessed with 16S rRNA gene terminal-restriction fragment length polymorphism and cloning, together with bacterial abundance and production measurements. In the early stages, open water and pancake ice were dominated by Alphaproteobacteria and Actinobacteria, which are common bacterial groups in Baltic Sea wintertime surface waters. The pancake ice bacterial communities were similar to the open-water communities, suggesting that the parent water determines the sea-ice bacterial community in the early stages of sea-ice formation. In consolidated young and thick ice, the bacterial communities were significantly different from water bacterial communities as well as from each other, indicating community development in Baltic Sea drift ice along with ice-type changes. The thick ice was dominated by typical sea-ice genera from classes Flavobacteria and Gammaproteobacteria, similar to those in polar sea-ice bacterial communities. Since the thick ice bacterial community was remarkably different from that of the parent seawater, results indicate that thick ice bacterial communities were recruited from the rarer members of the seawater bacterial community.

The structure of sea-ice bacterial communities is frequently different from that in seawater. Bacterial entrainment in sea ice has been studied with traditional microbiological, bacterial abundance, and bacterial production methods. However, the dynamics of the changes in bacterial communities during the transition from open water to frozen sea ice is largely unknown. Given previous evidence that the nutritional status of the parent water may affect bacterial communities during ice formation, bacterial succession was studied in under ice water and sea ice in two series of mesocosms: the first containing seawater from the North Sea and the second containing seawater enriched with algal-derived dissolved organic matter (DOM). The composition and dynamics of bacterial communities were investigated with terminal restriction fragment length polymorphism (T-RFLP), and cloning alongside bacterial production (thymidine and leucine uptake) and abundance measurements (measured by flow cytometry). Enriched and active sea-ice bacterial communities developed in ice formed in both unenriched and DOM-enriched seawater (0-6 days). I³-Proteobacteria dominated in the DOM-enriched samples, indicative of their capability for opportunistic growth in sea ice. The bacterial communities in the unenriched waters and ice consisted of the classes Flavobacteria, I±- and I³-Proteobacteria, which are frequently found in natural sea ice in polar regions. Furthermore, the results indicate that seawater bacterial communities are able to adapt rapidly to sudden environmental changes when facing considerable physicochemical stress such as the changes in temperature, salinity, nutrient status, and organic matter supply during ice formation.