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The Arctic sea-ice-scape is rapidly transforming. Increasing light penetration will initiate earlier seasonal primary production. This earlier growing season may be accompanied by an increase in ice algae and phytoplankton biomass, augmenting the emission of dimethylsulfide and capture of carbon dioxide. Secondary production may also increase on the shelves, although the loss of sea ice exacerbates the demise of sea-ice fauna, endemic fish and megafauna. Sea-ice loss may also deliver more methane to the atmosphere, but warmer ice may release fewer halogens, resulting in fewer ozone depletion events. The net changes in carbon drawdown are still highly uncertain. Despite large uncertainties in these assessments, we expect disruptive changes that warrant intensified long-term observations and modelling efforts.The Arctic is warming and undergoing rapid ice loss. This Perspective considers how changes in sea ice will impact the biogeochemistry and associated ecosystems of the region while calling for more observations to improve our understanding of this complex system.

Abstract 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°C and 4°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. Isolation, characterisation, host range and temperature adaptation of the first known Antarctic sea-ice virus isolates.

Sea ice is an important habitat for a wide variety of microalgal species. Depending on the species composition, sea ice can be a seeding source for pelagic phytoplankton blooms after ice melt in spring. Sea-ice algal communities were studied over 2 full winter seasons in 2014 and 2016 at Rothera Research Station, situated at the Western Antarctic Peninsula (WAP). Algal pigment patterns and microscopic observations were combined with photophysiological studies based on fluorescence analyses to monitor and explain the phenology of ice-algal species. Clear patterns in species succession were identified. Young sea ice contained a mixture of algal species including dinoflagellates, cryptophytes and diatoms like Chaetoceros spp. and Fragillariopsis spp. In winter, severe environmental conditions resulted in a decline in species diversity and selection towards heterotrophy. Pennate diatoms like Amphiprora kufferathii and Berkeleya adeliensis were the first to dominate the nutrient-enriched bottom-ice layers in early spring. The bottom communities exhibited a remarkably stable value for the photoadaptation parameter, Ek, of circa 25 µmol photons m–2 s–1. Whereas pennate diatoms were most abundant in spring ice, the initial seeding event linked to ice melt was associated with flagellate species. Haptophyte species like Phaeocystis antarctica and prymnesiophytes like Pyramimonas spp. best sustained the transition from sea ice to seawater. Comparison with previous studies shows that the seeding patterns observed in Ryder Bay were characteristic over the wider sea-ice domain, Arctic and Antarctic. Over the course of this century, the WAP is predicted to experience continuing thinning and decline in sea-ice cover. For the near future, we expect that especially microalgal communities of haptophytes and chlorophytes will benefit from the changes, with yet unknown implications for carbon fluxes and higher trophic levels.

Marine microbial communities undergo drastic changes during the seasonal cycle in high latitude seas. Despite the dominance of microbial biomass in the oceans, comprehensive studies on the seasonal changes of microbial plankton during the complete winter period are lacking. To study the seasonal variation in abundance of the microbial community, water samples were collected weekly in the Northern Baltic Sea from October to May. During ice cover from mid-January to April, samples from the sea ice and the underlying water were taken in addition to the water column samples. Abundances of bacteria, virus-like particles, nanoflagellates, and chlorophyll a concentrations were measured from sea ice, under-ice water, and the water column, and examined in relation to environmental conditions. All studied organisms had clear seasonal changes in abundance, and the sea-ice microbial community had an independent wintertime development compared to the water column. Bacteria were observed to have a key role in the biotic interactions in both ice and the water column, and the dormant period during the cold-water months (October–May) was limited to before ice formation. Our results provide the first insights into the temporal dynamics of bacteria and viruses during the whole cold-water season (October–May) in coastal high latitude seas, and demonstrate that changes in the environmental conditions are likely to affect bacterial dynamics and have implications on trophic interactions.

Sea-ice samples intended for biological analyses, e.g., chlorophyll- a , cell enumeration of algae and protozoa and primary production, are affected by the sampling and sample processing methods. In this study, we compared different sample processing methods by melting Baltic Sea ice samples in different ways (direct melting, buffered melting in filtered seawater (FSW) and buffered melting in artificial seawater at two different salinities with added nutrients) at two temperatures [+4 °C and room temperature (RT)]. We show that sea-ice samples intended for most commonly used biological analyses can be melted without the addition of FSW. In particular, adding artificial seawater should be avoided. To minimize biological processes, such as growth, death, predation and pigment degradation, the melting should be done rapidly at RT preferably by gently shaking the sample to keep the melt cool.

Ciliophora is a phylum of unicellular eukaryotes that are common and have pivotal roles in aquatic environments. Sea ice is a marine habitat, which is composed of a matrix of solid ice and pockets of saline water in which Ciliophora thrive. Here, we used phylogenetic placement to identify Ciliophora 18S ribosomal RNA reads obtained from wintertime water and sea ice, and assigned functions to the reads based on this taxonomic information. Based on our results, sea-ice Ciliophora assemblages are poorer in taxonomic and functional richness than under-ice water and water-column assemblages. Ciliophora diversity stayed stable throughout the ice-covered season both in sea ice and in water, although the assemblages changed during the course of our sampling. Under-ice water and the water column were distinctly predominated by planktonic orders Choreotrichida and Oligotrichida, which led to significantly lower taxonomic and functional evenness in water than in sea ice. In addition to planktonic Ciliophora, assemblages in sea ice included a set of moderately abundant surface-oriented species. Omnivory (feeding on bacteria and unicellular eukaryotes) was the most common feeding type but was not as predominant in sea ice as in water. Sea ice included cytotrophic (feeding on unicellular eukaryotes), bacterivorous and parasitic Ciliophora in addition to the predominant omnivorous Ciliophora. Potentially mixotrophic Ciliophora predominated the water column and heterotrophic Ciliophora sea ice. Our results highlight sea ice as an environment that creates a set of variable habitats, which may be threatened by the diminishing extent of sea ice due to changing climate.

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 (<4 μg l –1 ) chlorophyll- a (chl- a ) concentrations and consequent bacterial production. Typical sea-ice bacterial genera, for example, Octadecabacter , Polaribacter and Glaciecola , often abundant in spring and summer during the sea-ice algal bloom, predominated in the communities. The variability in bacterial community composition in the different ice types was mainly explained by the chl- a concentrations, suggesting that as in spring and summer sea ice, the sea-ice bacteria and algae may also be coupled during the Antarctic winter. Coupling between the bacterial community and sea-ice algae was further supported by significant correlations between bacterial abundance and production with chl- a . In addition, sulphate-reducing bacteria (for example, Desulforhopalus ) together with odour of H 2 S were observed in thick, apparently anoxic ice, suggesting that the development of the anaerobic bacterial community may occur in sea ice under suitable conditions. In all, the results show that bacterial community in Antarctic sea ice can stay active throughout the winter period and thus possible future warming of sea ice and consequent increase in bacterial production may lead to changes in bacteria-mediated processes in the Antarctic sea-ice zone.

The Baltic Sea is one of the world’s largest brackish water basins and is traditionally considered to be species poor. Here, we assessed the diversity of the nano-sized eukaryotic microbial wintertime community, using molecular ecological methods based on sequencing of small-subunit ribosomal RNA gene clone libraries. The results demonstrate that a rich community of small eukaryotes inhabits the Baltic Sea ice and water during winter. The community was dominated by alveolates and stramenopiles. Ciliates and cercozoans were the richest groups present, while in contrast to previous studies, diatoms showed a lower richness than expected. Furthermore, fungi and parasitic Syndiniales were present both in the water and in the sea ice. Some of the organisms in the sea-ice community were active, based on the RNA data, but a number of organisms were inactive or remnants from the freezing process. The results demonstrate that the sea-ice communities in the Baltic Sea are highly diverse and that water and ice of different ages include different protistan assemblages. Our study emphasizes the potential loss in biodiversity through diminishing ice cover as a result of climate change.

The effects of increased photosynthetically active radiation (PAR) and ultraviolet radiation (UVR) on species diversity, biomass and photosynthetic activity were studied in fast ice algal communities. The experimental set-up consisted of nine 1.44 m2 squares with three treatments: untreated with natural snow cover (UNT), snow-free (PAR + UVR) and snow-free ice covered with a UV screen (PAR). The total algal biomass, dominated by diatoms and dinoflagellates, increased in all treatments during the experiment. However, the smaller biomass growth in the top 10-cm layer of the PAR + UVR treatment compared with the PAR treatment indicated the negative effect of UVR. Scrippsiella complex (mainly Scrippsiella hangoei, Biecheleria baltica and Gymnodinium corollarium) showed UV sensitivity in the top 5-cm layer, whereas Heterocapsa arctica ssp. frigida and green algae showed sensitivity to both PAR and UVR. The photosynthetic activity was highest in the top 5-cm layer of the PAR treatment, where the biomass of the pennate diatom Nitzschia frigida increased, indicating the UV sensitivity of this species. This study shows that UVR is one of the controlling factors of algal communities in Baltic Sea ice, and that increased availability of PAR together with UVR exclusion can cause changes in algal biomass, photosynthetic activity and community composition. UVR in sea ice regulates algal biomass, vertical distribution, species composition and their photosynthetic activity; therefore, subsurface layers beneath the reach of UVR provide suitable growth conditions for sea ice algae.

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. Clear changes in the bacterial communities were observed in Baltic Sea drift-ice samples from open water to thick ice, leading to bacterial communities similar to those of polar sea ice.