Janne-Markus Rintala

Comparison of wintertime eukaryotic community from sea ice and open water in the Baltic Sea, based on sequencing of the 18S rRNA gene

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.

Role of Sea-ice Biota in Nutrient and Organic Material Cycles in the Northern Baltic Sea

Abstract This paper compiles biological and chemical sea-ice data from three areas of the Baltic Sea: the Bothnian Bay (Hailuoto, Finland), the Bothnian Sea (Norrby, Sweden), and the Gulf of Finland (Tvärminne, Finland). The data consist mainly of field measurements and experiments conducted during the BIREME project from 2003 to 2006, supplemented with relevant published data. Our main focus was to analyze whether the biological activity in Baltic Sea sea-ice shows clear regional variability. Sea-ice in the Bothnian Bay has low chlorophyll a concentrations, and the bacterial turnover rates are low. However, we have sampled mainly land-fast level first-year sea-ice and apparently missed the most active biological system, which may reside in deformed ice (such as ice ridges). Our limited data set shows high concentrations of algae in keel blocks and keel block interstitial water under the consolidated layer of the pressure ridges in the northernmost part of the Baltic Sea. In land-fast level sea-ice in the Bothnian Sea and the Gulf of Finland, the lowermost layer appears to be the center of biological activity, though elevated biomasses can also be found occasionally in the top and interior parts of the ice. Ice algae are light limited during periods of snow cover, and phosphate is generally the limiting nutrient for ice bottom algae. Bacterial growth is evidently controlled by the production of labile dissolved organic matter by algae because low growth rates were recorded in the Bothnian Bay with high concentrations of allochthonous dissolved organic matter. Bacterial communities in the Bothnian Sea and the Gulf of Finland show high turnover rates, and activities comparable with those of open water communities during plankton blooms, which implies that sea-ice bacterial communities have high capacity to process matter during the winter period.

Ice formation and growth shape bacterial community structure in Baltic Sea drift ice

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 extensive bloom of alternate-stage Prymnesium polylepis (Haptophyta) in the Baltic Sea during autumn–spring 2007–2008

During autumn 2007, an unusual increase in an algal species belonging to the order Prymnesiales was observed throughout the Baltic Sea Proper during routine national monitoring. Electron microscopical examination of the blooming species showed two types of flat scales – small and large – that resembled those of the alternate stage of Prymnesium polylepis. No spine-bearing scales were found. The 18S rDNA sequence data (n = 20, c. 1500 bp) verified the species identification as P. polylepis. There was up to 0.5% (7 bp) variability in the P. polylepis partial 18 S rDNA sequences from the Baltic Sea. These environmental sequences differed by 0–0.35% (0–4 bp) from cultured P. polylepis (isolate UIO036), and by 1.0–3.7% from other available Prymnesium sequences. The number of cells assumed to be P. polylepis began to increase in October 2007 coincidently with significantly calm and dry weather, and at their maximum the cells accounted for over 80% of the total phytoplankton biovolume in December–January. During February–April 2008, 95% of the Prymnesiales cells were in the size class of P. polylepis (>6 µm). The species attained bloom concentrations (>1 × 106 cells l–1) from March to May 2008. The species was observed throughout the Baltic Sea, except the Bothnian Bay, Gulf of Riga and the Kattegat. No toxic effects of the bloom were observed.

Physical and biological controls on DMS,P dynamics in ice shelf-influenced fast ice during a winter-spring and a spring-summer transitions

We report the seasonal and vertical variations of dimethylsulfide (DMS) and its precursor dimethylsulfoniopropionate (DMSP) in fast ice at Cape Evans, McMurdo Sound (Antarctica) during the spring-summer transition in 2011 and winter-spring transition in 2012. We compare the variations of DMS,P observed to the seasonal evolution of the ice algal biomass and of the physical properties of the ice cover, with emphasis on the ice texture and brine dynamics. Isolated DMS and DMSP maxima were found during both seasonal episodes in interior ice and corresponded to the occurrence of platelet crystals in the ice texture. We show that platelet crystals formation corresponded in time and depth to the incorporation of dinoflagellates (strong DMSP producers) in the ice cover. We also show that platelet crystals could modify the environmental stresses on algal cells and perturb the vertical redistribution of DMS,P concentrations. We show that during the winter-spring transition in 2012, the DMS,P profiles were strongly influenced by the development and decline of a diatom-dominated bloom in the bottom ice, with DMSP variations remarkably following chl a variations. During the spring-summer transition in 2011, the increase in brine volume fraction (influencing ice permeability) on warming was shown to trigger (1) an important release of DMS to the under-ice water through brine convection and (2) a vertical redistribution of DMSP across the ice.

Rhinomonas nottbecki n. sp. (Cryptomonadales) and Molecular Phylogeny of the Family Pyrenomonadaceae

The cryptomonad Rhinomonas nottbecki n. sp., isolated from the Baltic Sea, is described from live and fixed cells studied by light, scanning, and transmission electron microscopy together with sequences of the partial nucleus‐ and nucleomorph‐encoded 18S rRNA genes as well as the nucleus‐encoded ITS1, 5.8S, ITS2, and the 5′‐end of the 28S rRNA gene regions. The sequence analyses include comparison with 43 strains from the family Pyrenomonadaceae. Rhinomonas nottbecki cells are dorsoventrally flattened, obloid in shape; 10.0–17.2 μm long, 5.5–8.1 μm thick, and 4.4–8.8 μm wide. The inner periplast has roughly hexagonal plates. Rhinomonas nottbecki cells resemble those of Rhinomonas reticulata, but the nucleomorph 18S rRNA gene of R. nottbecki differs by 2% from that of R. reticulata, while the ITS region by 11%. The intraspecific variability in the ITS region of R. nottbecki is 5%. In addition, the predicted ITS2 secondary structures are different in R. nottbecki and R. reticulata. The family Pyrenomonadaceae includes three clades: Clade A, Clade B, and Clade C. All Rhinomonas sequences branched within the Clade C, while the genus Rhodomonas is paraphyletic. The analyses suggest that the genus Storeatula is an alternating morphotype of the genera Rhinomonas and Rhodomonas and that the family Pyrenomonadaceae includes some species that were described multiple times, as well as novel species.

An active bacterial community linked to high chl-a concentrations in Antarctic winter-pack ice and evidence for the development of an anaerobic sea-ice bacterial community

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 H2S 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.

Life associated with Baltic 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.

Sea-ice eukaryotes of the Gulf of Finland, Baltic Sea, and evidence for herbivory on weakly shade-adapted ice algae

To determine community composition and physiological status of early spring sea-ice organisms, we collected sea-ice, slush and under-ice water samples from the Baltic Sea. We combined light microscopy, HPLC pigment analysis and pyrosequencing, and related the biomass and physiological status of sea-ice algae with the protistan community composition in a new way in the area. In terms of biomass, centric diatoms including a distinct Melosira arctica bloom in the upper intermediate section of the fast ice, dinoflagellates, euglenoids and the cyanobacterium Aphanizomenon sp. predominated in the sea-ice sections and unidentified flagellates in the slush. Based on pigment analyses, the ice-algal communities showed no adjusted photosynthetic pigment pools throughout the sea ice, and the bottom-ice communities were not shade-adapted. The sea ice included more characteristic phototrophic taxa (49%) than did slush (18%) and under-ice water (37%). Cercozoans and ciliates were the richest taxon groups, and the differences among the communities arose mainly from the various phagotrophic protistan taxa inhabiting the communities. The presence of pheophytin a coincided with an elevated ciliate biomass and read abundance in the drift ice and with a high Eurytemora affinis read abundance in the pack ice, indicating that ciliates and Eurytemora affinis were grazing on algae.

Biogeochemical Impact of Snow Cover and Cyclonic Intrusions on the Winter Weddell Sea Ice Pack

Sea ice is a dynamic biogeochemical reactor and a double interface actively interacting with both the atmosphere and the ocean. However, proper understanding of its annual impact on exchanges, and therefore potentially on the climate, notably suffer from the paucity of autumnal and winter data sets. Here we present the results of physical and biogeochemical investigations on winter Antarctic pack ice in the Weddell Sea (R.V. Polarstern AWECS cruise, July-August 2013) which are compared with those from two similar studies conducted in the area in 1986 and 1992. The winter 2013 was characterized by a warm sea ice cover due to the combined effects of deep snow and frequent warm cyclones events penetrating southwards from the open Southern Ocean. These conditions were favorable to high ice permeability and cyclic events of brine movements within the sea ice cover (brine tubes), favoring relatively high chlorophyll-a (Chl-a) concentrations. We discuss the timing of this algal activity showing that arguments can be presented in favor of continued activity during the winter due to the specific physical conditions. Large-scale sea ice model simulations also suggest a context of increasingly deep snow, warm ice and large brine fractions across the three observational years, despite the fact that the model is forced with a snowfall climatology. This lends support to the claim that more severe Antarctic sea ice conditions, characterized by a longer ice season, thicker and more concentrated ice are sufficient to increase the snow depth and, somehow counter-intuitively, to warm the ice.