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Bacterial, fungal, and algal communities that colonize aquatic systems on glacial ice surfaces mediate biogeochemical reactions that alter meltwater composition and affect meltwater production and storage. In this study, we sought to improve understanding of microbial communities inhabiting the shallow aquifer that forms seasonally within the ice surface of a glacier's ablation zone (i.e., the weathering crust aquifer). Using a metagenomic approach, we compared gene contents of microbial assemblages in the weathering crust aquifer (WCA) of the Matanuska Glacier (Alaska, USA) to those recovered from supraglacial features and englacial ice. High abundances of Pseudomonadota, Cyanobacteriota, Actinomycetota, and Bacteroidota were observed across all samples, while taxa in class Gammaproteobacteria were found at significantly higher abundances in the weathering crust aquifer. The weathering crust aquifer samples also contained higher abundances of Dothideomycetes and Microbotryomyetes; fungal classes commonly observed in snow and other icy ecosystems. Phylogenetic analysis of 18S rRNA and rbcL gene sequences indicated high abundances of algae in the WCA that are closely related (> 98% and > 93% identity, respectively) to taxa of Ancylonema (Streptophyta) and Ochromonas (Ochrophyta) reported from glacial ice surfaces in Svalbard and Antarctic sea ice. Many functional gene categories (e.g., homeostasis, cellular regulation, and stress responses) were enriched in samples from the weathering crust aquifer compared to those from proximal englacial and supraglacial habitats, providing evidence for ecological specialization in the communities. The identification of phagotrophic phytoflagellate taxa and genes involved in mixotrophy implies that combined phototrophic and heterotrophic production may assist with persistence in the low light, low energy, and ephemeral conditions of the weathering crust environment. The compositional and functional differences we have documented indicate distinct microbial distributions and functional processes occur in the weathering crust aquifer environment, and we discuss how deciphering these nuances is essential for developing a more complete understanding of ecosystem biogeochemistry in supraglacial hydrological systems. [ABSTRACT FROM AUTHOR]

This study investigated the changes in sea ice temperature, microalgae species distribution, shape changes, and photosynthetic activity observed in the first‐year ice that forms in winter in Saroma‐ko Lagoon, Hokkaido, Japan. Temperatures at the bottom of the ice remained constant at −1.7°C, near the freezing point, while they varied between −6 and −1°C with diel fluctuations at the surface layer. Carefully collected algal samples showed high photosynthetic quantum yield and acclimation to the light intensities of individual ice layers; this indicates that the algal photosynthetic activity responds to dynamic changes in the ice environment, such as variations in temperature, salinity, and brine space. The algal communities consisted of more than 95% diatoms. Smaller algal cells were distributed in the upper layer of the sea ice compared to the lower layers. Chaetoceros sp., the dominant small‐cell species, was evenly distributed throughout the layers. In contrast, Detonula confervacea, the dominant large‐cell species, was unevenly distributed in the lower layer, with smaller colony size and cell volume in the upper layer. The shape differences observed in this species were thought to be a response to the changing environment within the first‐year sea ice. [ABSTRACT FROM AUTHOR]

The polymer-facilitated flux of ice algae on Arctic shelves can initiate benthic activity and growth after the nutritionally constrained winter period. Lipid-rich ice algae are readily consumed by benthos and those entering the sediment can benefit deposit feeders. Ice algae assimilated by benthic organisms cascade up multiple trophic levels within the benthic sub-web, re-entering the pelagic sub web through habitat coupling species. Pelagic predators can have significant ice-algal carbon signals obtained from the benthic compartment. Sympagic-pelagic-benthic coupling on Arctic shelves is expected to weaken with ongoing sea-ice change. This review discusses the phenology, quantity, and quality of ice-algal contributions to coupling, linked to thinning snow and ice cover including multi-year ice replacement. Predicting future coupling between marine sub-webs requires focused research that considers trophic markers of multiple carbon sources. [ABSTRACT FROM AUTHOR]

As Arctic sea ice and its overlying snow cover thin, more light penetrates into the ice and upper ocean, shifting the phenology of algal growth within the bottom of sea ice, with cascading impacts on higher trophic levels of the Arctic marine ecosystem. While field data or autonomous observatories provide direct measurements of the coupled sea ice‐algal system, they are limited in space and time. Satellite observations of key sea ice variables that control the amount of light penetrating through sea ice offer the possibility to map the under‐ice light field across the entire Arctic basin. This study provides the first satellite‐based estimates of potential sea ice‐associated algal bloom onset dates since the launch of CryoSat‐2 and explores how a changing snowpack may have shifted bloom onset timings over the last four decades. Plain Language Summary: Light plays a key role in sustaining life in the ocean. In the polar regions, sea ice and its overlying snow cover limit the amount of light reaching the upper ocean. Ice algae mainly live at the bottom centimeters of the ice and are important primary producers, using CO2 and inorganic nutrients to release O2 and create organic matter that is available to other trophic levels of the marine ecosystem. Every year rapid growth of bottom‐ice algae, known as the bloom onset, is triggered by the return of the sun and the increase in light intensities above that needed for photosynthesis to occur. Studies have shown that ice algae are shade adapted and can bloom under very low light conditions, yet the impact of recent changes in the snow‐ice cover that allow more light to penetrate earlier in the spring season remain uncertain. This study takes satellite data to map how ice algal bloom onset may be changing over time. Key Points: New processing methods to satellite data allow for daily under‐ice light mappingSatellite approaches allow for scaling up to the pan‐Arctic scalePhenology of ice‐algal bloom onset is strongly tied to snow cover variability [ABSTRACT FROM AUTHOR]

The polymer-facilitated flux of ice algae on Arctic shelves can initiate benthic activity and growth after the nutritionally constrained winter period. Lipid-rich ice algae are readily consumed by benthos and those entering the sediment can benefit deposit feeders. Ice algae assimilated by benthic organisms cascade up multiple trophic levels within the benthic sub-web, re-entering the pelagic sub web through habitat coupling species. Pelagic predators can have significant ice-algal carbon signals obtained from the benthic compartment. Sympagic-pelagic-benthic coupling on Arctic shelves is expected to weaken with ongoing sea-ice change. This review discusses the phenology, quantity, and quality of ice-algal contributions to coupling, linked to thinning snow and ice cover including multi-year ice replacement. Predicting future coupling between marine sub-webs requires focused research that considers trophic markers of multiple carbon sources.

Microalgae are the main source of the omega‐3 fatty acids eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA), essential for the healthy development of most marine and terrestrial fauna including humans. Inverse correlations of algal EPA and DHA proportions (% of total fatty acids) with temperature have led to suggestions of a warming‐induced decline in the global production of these biomolecules and an enhanced importance of high latitude organisms for their provision. The cold Arctic Ocean is a potential hotspot of EPA and DHA production, but consequences of global warming are unknown. Here, we combine a full‐seasonal EPA and DHA dataset from the Central Arctic Ocean (CAO), with results from 13 previous field studies and 32 cultured algal strains to examine five potential climate change effects; ice algae loss, community shifts, increase in light, nutrients, and temperature. The algal EPA and DHA proportions were lower in the ice‐covered CAO than in warmer peripheral shelf seas, which indicates that the paradigm of an inverse correlation of EPA and DHA proportions with temperature may not hold in the Arctic. We found no systematic differences in the summed EPA and DHA proportions of sea ice versus pelagic algae, and in diatoms versus non‐diatoms. Overall, the algal EPA and DHA proportions varied up to four‐fold seasonally and 10‐fold regionally, pointing to strong light and nutrient limitations in the CAO. Where these limitations ease in a warming Arctic, EPA and DHA proportions are likely to increase alongside increasing primary production, with nutritional benefits for a non‐ice‐associated food web. [ABSTRACT FROM AUTHOR]

In the Baltic Sea, cold-water adapted dinoflagellates and diatoms dominate the phytoplankton spring bloom of the Northern and Eastern Basins of the Baltic Sea. In the Central and Southern parts, where such species are less prominent, they cause occasional biomass peaks. We hypothesized that these dynamics correlate with ice cover, sea surface temperature (SST), and water transport processes, as the large Basins of the Central Baltic Sea are too deep to build-up blooms from their own seed banks. Long-term monitoring data from the past 40 and 20 years in the central and southern Baltic Sea, respectively, were analyzed here for biomass development of five cold-adapted taxa: the diatoms Pauliella taeniata, Thalassiosira baltica, Thalassiosira levanderi and Melosira spp. and the dinoflagellate Peridinella catenata. Results show that diatoms generally reached high biomass peaks in the 1980s and in shorter periods from 1995-1997, 2003- 2006, and 2010-2013 in all areas. We detected good correlations with the length of the ice cover period as well as low minimum and mean winter and spring SSTs. In contrast, biomass dynamics of the dinoflagellate P. catenata are more independent from these factors but have decreased strongly since the beginning of the 21st century. A numerical ocean model analysis confirmed the hypothesis that large blooms in the deep basins are seeded through water transport from adjacent shallow, ice-covered coastal areas such as the Gulf of Finland and the Gulf of Riga. Our results show that under ongoing climate warming, the common cold-water species may disappear from spring blooms in southern and central areas with unknown consequences for the ecosystem.

Melting sea ice is a seasonal source of iron (Fe) to the Southern Ocean (SO), where Fe levels in surface waters are otherwise generally too low to support phytoplankton growth. However, the effectiveness of sea-ice Fe fertilization in stimulating SO primary production is unknown since no data exist on Fe uptake by microorganisms in the sea-ice environment. This study reports a unique dataset on Fe uptake rates, Fe-to-carbon (C) uptake ratio (Fe uptake normalized to C uptake) and Fe:C uptake rate (Fe uptake normalized to biomass) by in situ microbial communities inhabiting sea ice and the underlying seawater. Radioisotopes 55Fe and 14C were used in short-term uptake experiments during the 32-day Ice Station POLarstern (ISPOL) time series to evaluate the contributions of small (0.8–10 µm) and large (> 10 µm) microbes to Fe uptake. Overall, results show that over 90% of Fe was bound to the outside of the cells. Intracellular Fe (Feintra) uptake rates reached up to 68, 194, and 203 pmol Fe L−1d− 1 in under-ice seawater, bottom ice, and top ice, respectively. Inorganic carbon uptake ranged between 0.03 and 3.2 µmol C L−1 d−1, with the lowest rate observed in under-ice seawater. Importantly, between the start and end of ISPOL, we observed a 30-fold increase in Feintra normalized to carbon biomass in bottom sea ice. This trend was likely due to changes in the microbial community from a dominance of large diatoms at the start of the survey to small diatoms later in the season. As the Antarctic icescape and associated ecosystems are changing, this dataset will help inform the parameterisation of sea-ice biogeochemical and ecological models in ice-covered regions. [ABSTRACT FROM AUTHOR]

Anthropogenic climate change is causing changes to the Arctic sea-ice system with implications for the magnitude and timing of Arctic pelagic and ice-associated (sympagic) primary production that influences food web interactions. Ringed seals (Pusa hispida) are generalist predators that, as a species experience vastly different icescapes from low to high-Arctic latitudes. Quantifying spatial variation in their diet can help us understand how changes in sea-ice dynamics affect trophic interactions in Arctic marine food webs. However, multiple complementary analytical tools to examine variation in carbon source use and trophic dynamics in the diet of ringed seals have not yet been applied across their latitudinal range in the Arctic. We conducted stable isotope analysis (δ13C and δ15N) and measured highly branched isoprenoid diatom lipid biomarkers of ringed seals from the low, intermediate, and high Arctic (from 61.1°N to 77.5°N) to investigate spatial variation in their carbon source use and trophic position in relation to sea-ice dynamics. Both δ13C and highly branched isoprenoids indicated that ringed seals from higher latitudes had more sympagic carbon in their diet (liver δ13C: −18.3 ± 0.2 ‰, HBI: 89.9 ± 2.08 %) than ringed seals at lower latitudes (liver δ13C: −21.1 ± 0.1 ‰, HBI: 22.0 ± 2.73 %). Ringed seal trophic position increased from the low (3.78 ± 0.02) to high (4.76 ± 0.03) Arctic, suggesting increased fish consumption or a different trophic structure coinciding with the latitudinal change in carbon source. Ringed seals demonstrated a clear shift from low to high Arctic in the relative contribution of phytoplanktonic vs sympagic primary production. These patterns are likely linked to the vastly different icescapes in these environments and demonstrate that shifts in primary producer composition and Arctic food webs can be identified in ringed seal diets. Information on these prey and energy shifts over large spatial scales also provides insights into potential future changes to Arctic ecosystem function with continued sea-ice decline.

In the seasonal ice zone (SIZ), sea-ice algae have been hypothesized to influence phytoplankton species composition in seawater after melting from sea ice. However, information on ice-algal species composition during summer (December–February) was small particularly in the Indian sector of the Southern Ocean. We investigated sea-ice macronutrient concentrations and relationships between phytoplankton and ice-algal species composition in this region during January of 2018–2020. The species composition of ice algae each year (after weighted-averaged by sample volume for each sea ice) was dominated by the diatom Fragilariopsis cylindrus (78–89% of total cells). This result differed from ice-algal compositions in spring but consistent with those in summer, indicating seasonal changes in species composition of ice algae at East Antarctic SIZ. Using all years' sea-ice data, we found nitrate + nitrite: phosphate ratio declined significantly with increased algal concentrations. Diatoms' uptake ratio of macronutrients and remineralization of phosphorus from their products (e.g., dissolved organic matter) may result in this relationship, which indicates algal accumulation especially of diatoms in sea ice. At the ice-sampling stations, phytoplankton was dominated by F. cylindrus, while at the northern stations, higher proportions of F. kerguelensis, Pseudo-nitzschia prolongatoides/subcurvata, and Gymnodiniales were observed. The similarity index of sea-ice and phytoplankton composition was negatively correlated with days elapsed since sea-ice retreat and surface seawater temperature at each sampled station. This suggests that phytoplankton species composition might be influenced by release of ice algae where the influence of sea ice is strong during summer. [ABSTRACT FROM AUTHOR]

Novel observations of the seasonal evolution of an ice algal bloom on the Chukchi shelf were collected by two autonomous buoys deployed 180 km apart in first‐year drifting sea ice. High attenuation of blue light in the bottom of the ice indicated considerable accumulation of ice algae biomass with derived Chlorophyll‐a concentrations (Chl a) up to 184 mg m−2. Differences in the magnitude and persistence of ice algae biomass under each buoy appear to have been driven by differences in snow thickness, as ice thickness was similar between the sites. Minimal snow cover (0.02 m) around one buoy was associated with algae growth beginning in mid‐May and lasting 70 days. The second buoy had notably more snow (0.4 m), causing ice algae production to lag behind the first site by approximately 4 weeks. The delay in growth diminished the peak of ice algae Chl a and duration compared to the first site. Light attenuation through the ice was intense enough at both buoys to have a potentially inhibiting impact on water column phytoplankton Chl a. Modeling ice algae growth with observed light intensities determined that nutrients were the limiting resource at the low snow site. In contrast, the algae at the high snow site were light‐limited and never nutrient‐limited. These data point toward changes in ice algae phenology with an earlier and longer window for growth; and nutrients rather than light determining the longevity and magnitude of production. Plain Language Summary: Ice algae growing on the underside of Arctic sea ice require light and nutrients. Light levels are enhanced beneath sea ice that is thinner and has less snow accumulation, creating a favorable environment for algal growth. This study describes results from two autonomous instrument systems installed in Arctic sea ice in March 2018. Sensors measured the amount of light transmitted through the ice and snow. Changes in the light were connected to changes in the magnitude of algae present. One site had 0.02 m of snow, and the other had 0.4 m; ice thickness was similar between the sites. Ice algae appeared in mid‐May at the low‐snow site, growth at the high‐snow site began 4 weeks later. Numerical modeling showed that algae at the low snow site had abundant light, with growth limited by nutrients. In comparison, light was the limiting resource at the high snow site. Additionally, absorption of light by ice algae reduced the growth of water column phytoplankton under the ice. These data show that as snow cover is reduced on sea ice, ice algae could start growing earlier in the season and reach higher concentrations. Key Points: Autonomous buoys captured the seasonal evolution of ice algae growth in first‐year Arctic sea ice, bloom initiation determined by snow depthEarly bloom onset and 70‐day growth window were observed, evidence that shifts in phenology projected by climate models have occurredNutrients limited ice algae growth; movement of the ice promoted turbulent diffusion of nutrients supporting high biomass [ABSTRACT FROM AUTHOR]

The article presents data on the composition and abundance of algal flora in the coastal stratified Kislo-Sladkoe Lake (the Karelian coast of the White Sea) after its flushing with seawater in autumn. The seasonal dynamics and vertical structure of the phytoplankton were monitored from March 17 to October 3, 2019. We also studied ice flora on March 17 and April 1. The largest carbon biomass of ice algae (0.79 g C/m3) dominated by Nitzschia frigida was recorded in April. The integrated biomass of phytoplankton in the water column varied from 0.01 g C/m2 in March to 5.83 g C/m2 during spring bloom in late May (when it was dominated by Chaetoceros invisibilis) and remained at the level of 0.07 to 0.29 g C/m2 until October; heterotrophic forms replaced autotrophic algae during this period. A regression analysis has shown that the structure of the phytoplankton was influenced by length of daylight, water temperature, dissolved oxygen level, depth, pH, salinity, and illuminance. [ABSTRACT FROM AUTHOR]

The Chukchi Sea is the inflow shelf between the Arctic and the Pacific. A comprehensive research on algal dynamics, physical–biological interactions, and impacts on nutrient drawdown was conducted in summer 2010. The study found a phytoplankton bloom with chlorophyll a (Chla) > 10 mg m−3 at ice edge in July. There were 36 shared algal species in both sea ice and open water, and those species dominated numerically both environments highlighting the seeding role of ice algae for the phytoplankton bloom. The succession pattern of diatom genera at three phases from sea-ice to open-water bloom and post-bloom conditions was revealed, and the phytoplankton abundance was positively correlated with the concentration of > 20 µm fractionated Chla. An anticyclonic eddy was detected in July and significantly reduced nutrient inventories in the impacted area relative to historical values. Phytoplankton composition was an important factor for both variable drawdown ratios of silicate to dissolved inorganic nitrogen (1.31–3.37) and dissolved inorganic nitrogen to dissolved inorganic phosphorus (7.62–12.32); however, both ratios also changed between July and August. The results indicated highly dynamic biogeochemical processes during the summer. In addition, the lower bound of silicate drawdown was comparable to the amount imported through the Bering Strait, indicating the Chukchi Sea is a significant silicate sink. [ABSTRACT FROM AUTHOR]

Arctic summer sea ice extent is decreasing and thinning, forming melt ponds that cover more than 50% of the sea ice area during the peak of the melting season. Despite of this, ice algal communities in melt ponds are understudied and so are their contribution to the Arctic Ocean primary production and carbon turnover. While melt ponds have been considered as low productive, recent studies suggest that accumulated ice algal potentially facilitate high and yet overlooked rates of carbon turnover. Here we report on ice algal communities forming dense mats not previously described, collected from melt ponds in the northern Barents Sea in July. We document on distinct layered and brown colored mats with high carbon assimilation and net primary production rates compared to ice algal communities and aggregates, in fact comparable to benthic microalgae at temperate tidal flats. Rates of gross and net primary production, as well as community respiration rates were obtained from oxygen micro profiling, and carbon assimilation calculations were supported by 14C incubations, pigment analysis and light microscopy examinations. The melt pond algal mats consisted of distinct colored layers and differed from aggregates with a consisted layered structure. We accordingly propose the term melt pond algal mats, and further speculate that these dense ice algal mats may provide an important yet overlooked source of organic carbon in the Arctic food-web. A foodweb component likely very sensitive to climate driven changes in the Arctic Ocean and pan-Arctic seas.

The impact of the rapidly changing Arctic on zooplankton community structure and seasonal behaviour is not yet understood. Here we examine 6 months of under-ice zooplankton observations from the N-ICE2015 expedition (January to June 2015) in the Nansen Basin and on the Yermak Plateau north of Svalbard. Stratified sampling in the water column was done with MultiNet during the entire expedition, and sampling in the upper 5 m below sea ice was performed during April-May by divers using a hand-held net. Hydrographic conditions were dominated by northward-flowing warm and saline Atlantic Water at intermediate depth, and southward-flowing cold Polar Surface Water in the upper 100 m. The mesozooplankton was dominated by copepods. Most numerous were the small ubiquitous Oithona similis in the upper 200 m, with Microcalanus spp. and Triconia borealis further down the water column. Calanus finmarchicus dominated among the Calanus species while Metridia longa was also numerous. The most abundant deep-water copepods were Paraeuchaeta spp. and Spinocalanus spp. Arrow worms (Chaetognatha) and comb jellies (Ctenophora) were the most numerous non-copepods. The mesozooplankton community was more dependent on surrounding water mass characteristics, such as salinity and depth, than geographical location. Algal food availability, which was closely linked to seasonality, explained the community changes seen in surface waters in May and June due to seasonal ascent and recruitment. Seasonal changes from winter to spring mostly involved an increase in the herbivorous C. finmarchicus and its nauplii in the upper 200 m of the water column coinciding with the peak of the phytoplankton bloom in late May. The Yermak Plateau and adjacent Nansen Basin were characterised by oceanic North Atlantic and Arctic species, many of which are deep water specialists. Despite the late onset of the spring bloom due to consolidated sea ice, both North Atlantic and Arctic species successfully reproduced in the study area. This explains the species-rich mesozooplankton community in this region as opposed to the less productive central Arctic Ocean. Future prospects of less sea ice and earlier onset of the bloom will likely be positive for the overall secondary production by both Arctic and boreal zooplankton in this region.

Oil production in Arctic ice-covered areas poses a risk for pollution of the ecosystem including that within the brine channel network of sea ice. Sea-ice autotrophs contribute substantially to Arctic primary production, but are inherently difficult to test for oil exposure responses in situ. This study had two objectives, first, we developed a suitable lab-based mesocosm system, second, we tested oil effects on sea-ice algae. Specifically, we investigated if Alaska North Slope crude oil exposure reduces ice algal abundance, biomass and concentration of extracellular polymeric substances (EPS) using indoor ice tanks over a 10-day exposure period. Six tanks in one cold room were used in pairs for the following treatments: (1) control, (2) oil release as a layer under ice and (3) release of dispersed oil. All tanks were inoculated with sea-ice microbial communities collected from Utqiaġvik, Alaska. After 10 days of exposure, the abundance of algae, dominated by the pennate diatom genus Nitzschia, and the concentrations of EPS and chlorophyll a were significantly lower in the oiled treatments compared to the control. We suggest light attenuation by the oil, reduced algal mobility, and oil toxicity as causes for this reduction. Observed changes in cell fluorescence characteristics based on DNA staining could be linked to the oil exposure and could provide a new tool for assessment of toxicity in microalgae. [ABSTRACT FROM AUTHOR]

In this study, we present unique data collected with a Surface and Under-Ice Trawl (SUIT) during five campaigns between 2012 and 2017, covering the spring to summer and autumn transition in the Arctic Ocean, and the seasons of winter and summer in the Southern Ocean. The SUIT was equipped with a sensor array from which we retrieved: sea-ice thickness, the light field at the underside of sea ice, chlorophyll a concentration in the ice (in-ice chl a), and the salinity, temperature, and chl a concentration of the under-ice water. With an average trawl distance of about 2 km, and a global transect length of more than 117 km in both polar regions, the present work represents the first multi-seasonal habitat characterization based on kilometer-scale profiles. The present data highlight regional and seasonal patterns in sea-ice properties in the Polar Ocean. Light transmittance through Arctic sea ice reached almost 100% in summer, when the ice was thinner and melt ponds spread over the ice surface. However, the daily integrated amount of light under sea ice was maximum in spring. Compared to the Arctic, Antarctic sea-ice was thinner, snow depth was thicker, and sea-ice properties were more uniform between seasons. Light transmittance was low in winter with maximum transmittance of 73%. Despite thicker snow depth, the overall under-ice light was considerably higher during Antarctic summer than during Arctic summer. Spatial autocorrelation analysis shows that Arctic sea ice was characterized by larger floes compared to the Antarctic. In both Polar regions, the patch size of the transmittance followed the spatial variability of sea-ice thickness. In-ice chl a in the Arctic Ocean remained below 0.39 mg chl a m−2, whereas it exceeded 7 mg chl a m−2 during Antarctic winter, when water chl a concentrations remained below 1.5 mg chl a m−2, thus highlighting its potential as an important carbon source for overwintering organisms. The data analyzed in this study can improve large-scale physical and ecosystem models, habitat mapping studies and time series analyzed in the context of climate change effects and marine management.

Phytoplankton communities in seasonally ice-covered areas are largely affected by ice algae. The Okhotsk Sea is the southernmost sea ice zone in the northern hemisphere with a sizeable seasonal ice cover, thus ice algae of the Okhotsk sea ice have a large potential to seed the early spring bloom. Little is known about the Okhotsk ice algal communities and their seeding effects. We investigated the dynamics of the composition and the photophysiological performances of an Okhotsk ice algal community in a 6-day laboratory incubation experiment that simulated the natural ice melt conditions. Centric diatoms, especially Thalassiosira spp., overwhelmingly dominated the ice algal community throughout incubation, whereas pennate diatoms, mostly Navicula and Nitzschia, showed little growth with much higher mortality. The maximum photochemical efficiency of Photosystem II (Fv/Fm) was the lowest at the beginning of the ice melt, suggesting a suppressed photosynthetic functioning by changes in salinity. The cellular pigment contents decreased by 30% due to osmotic stress, evidenced by deformed plastids under a microscope. The transcript level of the rbcL gene that encodes the large subunit of RubisCO was significantly higher during ice melt and decreased in the no-ice period, suggesting an urgent need for carbon fixation under the melt condition. Full recovery of photosynthesis and growth was also made after complete ice melt. Our results indicated high seeding potential of Thalassiosira to spring blooms owing to their photophysiological plasticity to dynamic salinity changes.

High concentrations of microplastics have been found in sea ice but the mechanisms by which they get captured into the ice and which role ice algae might play in this process remain unknown. Similarly, we do not know how the presence of microplastics might impact the colonization of sea ice by ice algae. To estimate the ecological impact of microplastics for Polar ecosystems, it is essential to understand their behaviour during ice formation and possible interactions with organisms inhabiting sea ice.In this study we tested the interaction between the ice algae Fragillariopsis cylindrus and microplastic beads with and without sea ice present and, in a third experiment, during the process of ice formation. With sea ice present, we found significantly less algae cells in the ice when incubated together with microplastics compared to the incubation without microplastics. However, during ice formation, the presence of microplastics did not impact the colonisation of the ice by F. cylindrus cells. Further, we observed a strong correlation between salinity and the relative amount of beads in the water and ice. With increasing salinity of the water, the relative amount of beads in the water decreased significantly. At the same time, the relative amount of beads in the ice increased significantly with increasing ice salinity. Both processes were not influenced by the presence of F. cylindrus. Also, we found indications that the presence of algae can affect the amount of microplastic beads sticking to the container walls. This could indicate that EPS produced by ice algae plays a significant role in surface binding properties of microplastics.Overall, our results highlight that the interactions between algae and microplastics have an influence on the uptake of microplastics into sea ice with possible implications for the sea ice food web.

The large declines in Arctic sea-ice age and extent over the last decades could have altered the diversity of sea-ice associated unicellular eukaryotes (referred to as sea-ice protists). A time series from the Russian ice-drift stations from the 1980s to the 2010s revealed changes in community composition and diversity of sea-ice protists from the Central Arctic Ocean. However, these observations have been biased by varying levels of taxonomic resolution and sampling effort, both of which were higher in the early years at drift stations on multiyear sea ice (MYI) in the Central Arctic Ocean. We here combine the Russian ice-drift station data with more recent data to (1) identify common sea-ice protists (in particular diatoms) in drifting sea ice of the Central Arctic Ocean; (2) characterize the potential change in such communities over 35 years in terms of species number and/or community structure; and (3) relate those shifts to relevant environmental factors. In terms of relative abundance, pennate diatoms were the most abundant sea-ice protists across the Arctic, contributing 60% on average of counted cells. Two pennate colony-forming diatom species, Nitzschia frigida and Fragilariopsis cylindrus, dominated at all times, but solitary diatom species were also frequently encountered, e.g., Cylindrotheca closterium and Navicula directa. Multiyear sea ice contained 39% more diatom species than first-year ice (FYI) and showed a relatively even distribution along entire sea-ice cores. The decrease in MYI over the last decades explained the previously reported decreases in sea-ice protist diversity. Our results also indicate that up to 75% of diatom species are incorporated into FYI from the surrounding sea ice and the water column within a few months after the initial formation of the ice, while the remaining 25% are incorporated during ice drift. Thus, changing freeze-up scenarios, as currently witnessed in the Central Arctic, might result in long-term changes of the biodiversity of sea-ice protists in this region.

Field observations suggest that oceanic emissions of dimethylsulfide (DMS) may play a dominant role in the production of Arctic aerosols and clouds and therefore modulate the surface irradiance, during spring and summer. DMS is produced not only in the water column but also in various sea ice habitats. The ongoing recession of Arctic sea ice is expected to enhance DMS emissions, but the magnitude of this increase is highly uncertain. Here we investigate the spatiotemporal variability in bottom ice and sea surface DMS concentrations and fluxes using a regional sea ice‐ocean physical‐biogeochemical model. Model results indicate that the observed accelerated decline of Arctic sea ice extent since the beginning of the 21st century is associated with upward trends in May–August pan‐Arctic‐averaged sea surface DMS concentration and sea‐to‐air DMS flux. On the other hand, strong interannual variability and statistically insignificant trends are found for bottom ice DMS concentration and ice‐to‐sea DMS flux, owing to the counteracting effects of the shrinking horizontal extent and the vertical thinning of sea ice on ice algal production. The pan‐Arctic DMS climatology products based on model simulation and satellite algorithms provide dynamically based spatial details that are absent in the in situ measurement‐based climatology due to limited spatiotemporal data coverage and inevitable extrapolation bias. Lastly, model results indicate that the bottom ice DMS and its precursor dimethylsulfoniopropionate production can be the only local source of oceanic DMS emissions into the atmosphere during May prior to pelagic blooms, suggesting that it may be a key component of the biological control on Arctic climate at that time. Key Points: Model results reveal a 39% increase in the May–August and pan‐Arctic mean sea‐to‐air DMS flux since the beginning of the 21st centuryIn contrast, bottom ice DMS shows no apparent trend but exhibits strong interannual variabilityBottom ice DMSP and DMS can become the sole source of sea surface DMS concentration and sea‐to‐air flux over extended regions in May [ABSTRACT FROM AUTHOR]

Arctic sea ice algae produce extracellular organic products, which, as bioconditioners of seawater, may stimulate early summer growth of pelagic, under-sea-ice phytoplankton in low light and low temperature conditions. Sea ice algae are inhibited or decline in numbers if prematurely exposed to high light conditions, thereby reducing their ability to produce bioconditioners. As climate change creates an early reduction or removal of snow and sea ice cover, the result may be a decrease in primary phytoplankton production. [ABSTRACT FROM AUTHOR]

This work summarizes ice algal studies, presented as biomass and species temporal and spatial distribution, during 11 cruises conducted between 1986 and 2012. The majority of the biomass was found as loosely attached sub-ice algal layers, and sampling required diving. A maximum of 40 mg chlorophyll m−2 and 15.4 × 109 cells m−2 was measured in May. The species diversity was separated in zones based on ice thickness, with the highest biodiversity in the medium-thick ice of 30–80 cm. Nitzschia frigida was the most common species. There was a significant positive relationship between the dominance of this species and ice thickness, and it dominated completely in thick ice. Other common species, such as N. promare and Fossulaphycus arcticus reacted oppositely, by becoming less dominant in thick ice, but the positive correlation between total cell numbers and number of these three species indicated that they would most likely dominate in most populations. Melosira arctica was found several times below medium-thick annual ice. Algae occurred from top to bottom in the ice floes and in infiltration layers, but in very low numbers inside the ice. The bipolar dinoflagellates Polarella glacialis inhabited the ice, both as vegetative cells and cysts. The algal layers detached from the ice and sank in late spring when melting started. The cells in the sediments form an important food source for benthic animals throughout the year. Fjord populations survive the winter on the bottom and probably form next year’s ice algal inoculum. A few ‘over-summer’ populations found in sheltered locations might provide supplementary food for ice amphipods in late summer. The future faith of the ice flora is discussed in view of a warmer climate, with increased melting of the Arctic ice cover.

Ice algae are a key component in polar marine food webs and have an active role in large-scale biogeochemical cycles. They remain extremely under-sampled due to the coarse nature of traditional point sampling methods compounded by the general logistical limitations of surveying in polar regions. This study provides a first assessment of hyperspectral imaging as an under-ice remote-sensing method to capture sea-ice algae biomass spatial variability at the ice/water interface. Ice-algal cultures were inoculated in a unique inverted sea-ice simulation tank at increasing concentrations over designated cylinder enclosures and sparsely across the ice/water interface. Hyperspectral images of the sea ice were acquired with a pushbroom sensor attaining 0.9 mm square pixel spatial resolution for three different spectral resolutions (1.7, 3.4, 6.7 nm). Image analysis revealed biomass distribution matching the inoculated chlorophyll a concentrations within each cylinder. While spectral resolutions >6 nm hindered biomass differentiation, 1.7 and 3.4 nm were able to resolve spatial variation in ice algal biomass implying a coherent sensor selection. The inverted ice tank provided a suitable sea-ice analogue platform for testing key parameters of the methodology. The results highlight the potential of hyperspectral imaging to capture sea-ice algal biomass variability at unprecedented scales in a non-invasive way.

Sea ice algae represent a key energy source for many organisms in polar food webs, but estimating their biomass at ecologically appropriate spatiotemporal scales remains a challenge. Attempts to extend ice-core derived biomass to broader scales using remote sensing approaches has largely focused on the use of under-ice spectral irradiance. Normalized difference index (NDI) based algorithms that relate the attenuation of irradiance by the snow-ice-algal ensemble at specific wavelengths to biomass have been used to explain up to 79% of the biomass of algae in limited areas. Application of these algorithms to datasets collected using tethered remotely operated vehicles (ROVs) has begun, generating methods for spatial sampling at scales and spatial resolution not achievable with ice-core sampling. Successful integration of radiometers with untethered autonomous underwater vehicles (AUVs) offers even greater capability to survey broader regions to explore the spatial heterogeneity of sea ice algal communities. This work describes the pilot use of an AUV fitted with a multispectral irradiance sensor to estimate ice-algal biomass along transects beneath land-fast sea ice (∼2 m thick with minimal snow cover) in McMurdo Sound, Antarctica. The AUV obtained continuous, repeatable, multi-band irradiance data, suitable for NDI-type approaches, over transects of 500 m, with an instrument footprint of 4 m in diameter. Algorithms were developed using local measurements of ice algae biomass and spectral attenuation of sea ice and were able to explain 40% of biomass variability. Relatively poor performance of the algorithms in predicting biomass limited the confidence that could be placed in biomass estimates from AUV data. This was attributed to the larger footprint size of the optical sensors integrating small-scale biomass variability more effectively than the ice core in the platelet-dominated ice algal habitat. Our results support continued development of remote-sensing of sea ice algal biomass at m–km spatial scales using optical methods, but caution that footprint sizes of calibration data (e.g., coring) must be compatible with optical sensors used. AUVs offer autonomous survey techniques that could be applied to better understand the horizontal variability of sea ice algae from nearshore ice out to the marginal ice zone.

Sea ice algae are an important contributor of primary production in the Arctic ecosystem. Within the bottom-ice environment, access to nutrients from the underlying ocean is a major factor controlling production, phenology, and taxonomic composition of ice algae. Previous studies have demonstrated that tides and currents play an important role in driving the flux of nutrients to bottom-ice algal communities when biological demand during the spring bloom is high. In this study we investigate how surface currents under landfast first-year ice influence nutrient supply based on stoichiometric composition, algal chlorophyll a biomass and species composition during spring 2016, in Dease Strait, Nunavut. Stronger water dynamics over a shoaled and constricted strait dominated by tidal currents (tidal strait) supported turbulent flow more than 85% of the deployment duration in comparison to outside the tidal strait in an embayment where turbulent flow was only evidenced a small percentage (

Recent studies have highlighted the importance of ice-algal blooms in driving darkening and therefore surface melt of the Greenland Ice Sheet (GrIS). However, the contribution of fungal and bacterial communities to this microbially driven albedo reduction remains unconstrained. To address this significant knowledge gap, fungi were isolated from key GrIS surface habitats (surface ice containing varying abundance of ice algae, supraglacial water, cryoconite holes, and snow), and a combination of cultivation and sequencing methods utilized to characterize the algal-associated fungal and bacterial diversity and abundance. Six hundred and ninety-seven taxa of fungi were obtained by amplicon sequencing and more than 200 fungal cultures belonging to 46 different species were isolated through cultivation approaches. Basidiomycota dominated in surface ice and water samples, and Ascomycota in snow samples. Amplicon sequencing revealed that bacteria were characterized by a higher diversity (883 taxa detected). Results from cultivation as well as ergosterol analyses suggested that surface ice dominated by ice algae and cryoconite holes supported the highest fungal biomass (104–105 CFU/100 ml) and that many fungal taxa recognized as endophytes and plant pathogens were associated with dark ice characterized by a high abundance of ice algae. This paper significantly advances this field of research by investigating for the first time the fungal abundance and diversity associated with algal blooms causing the darkening of the GrIS. There is a strong association between the abundance and diversity of fungal species and the blooming of algae on the surface ice of the Greenland Ice Sheet.

Snow cover on sea ice is the most important factor controlling light availability for sea ice algae, but it is predicted by climate models to become more variable and stochastic. Here, we document effects of a sudden, complete loss of the entire snow cover on first-year sea ice at Kangerlussuaq Fjord, West Greenland, due to a natural Föhn wind event that caused a ca. 17 °C air temperature increase over 36 h. We applied Imaging-PAM fluorometry to examine effects of snow cover on algal distribution and photobiology and observed a rapid decrease in algal biomass associated with loss of the skeletal ice crystal layer on the underside of the ice that had supported most of the visible algae. Furthermore, the remaining algae were photobiologically stressed, as seen in a significant decrease in the dark-acclimated fluorescence yield (ΦPSII_max) from 0.55 before snow loss to 0.41 after. However, recovery in the dark suggested that non-photosynthetic quenching was successfully dissipating excess energy in the community and that there was little photodamage. An observed decrease in the photosynthetic efficiency α from 0.22 to 0.16 µmol é m−2 s−1 is therefore likely to be due to photoacclimation and the change in community composition. Centric diatoms and flagellates were the main taxa lost in the snow loss event, whereas the sea ice specialist Nitzschia frigida increased in numbers. These observations are similar to those seen in artificial snow-clearing experiments and consistent with snow clearing being a useful approach for investigating the complex interactions between snow cover, irradiance fluctuations, and ice algal performance.

Seasonal, interannual, and decadal variations in the Arctic ice‐algal productivity for 1980–2009 are investigated using daily outputs from five sea ice‐ocean ecosystem models participating in the Forum for Arctic Modeling and Observational Synthesis project. The models show a shelf‐basin contrast in the spatial distribution of ice‐algal productivity (ice‐PP). The simulated ice‐PP substantially varies among the four subregions (Chukchi Sea, Canada Basin, Eurasian Basin, and Barents Sea) and among the five models, respectively. The simulated annual total ice‐PP has no common decadal trend at least for 1980–2009 among the five models in any of the four subregions, although the simulated snow depth and sea‐ice thickness in spring are mostly declining. The model intercomparison indicates that an appropriate balance of stable ice‐algal habitat (i.e., sea‐ice cover) and enough light availability is necessary to retain the ice‐PP. The multi‐model averages show that the ice‐algal bloom timing shifts to an earlier date and that the bloom duration shortens in the four subregions. However, both the positive and negative decadal trends in the timing and duration are simulated. This difference in trends are attributed to temporal shifts among different types of ice‐algal blooms: long‐massive, short‐massive, long‐gentle, and short‐gentle bloom. The selected value for the maximum growth rate of the ice‐algal photosynthesis term is a key source for the inter‐model spreads. Understanding the simulated uncertainties on the pan‐Arctic and decadal scales is expected to improve coupled sea ice‐ocean ecosystem models. This step will be a baseline for further modeling/field studies and future projections. Key Points: An appropriate balance of stable sea‐ice cover and enough light availability is necessary to retain ice‐algal productivityAnnual total ice‐algal productivity has no common decadal trend for 1980–2009 among the five marine ecosystem modelsThe multi‐model averages show that ice‐algal bloom timing shifts to an earlier date and the bloom duration shortens in four subregions [ABSTRACT FROM AUTHOR]

In the Bering Sea shelf, annual changes in the sea ice extent are large. In this study, we compare the viable diatom resting stages in sediments during the summer of 2017 when the sea ice retreat was late and 2018 when the sea ice retreat was early. South of St. Lawrence Island, the germinating cell number was 10–100 times greater in 2018 than it was in 2017. The taxonomic composition also showed large annual differences: Fragilariopsis/Fossula spp., which are ice algae species, were abundant in 2017, but Thalassiosira spp. dominated in 2018. Satellite observations confirmed that sea ice diminished before the ice-edge bloom in 2018, but sea ice remained until the ice-edge bloom in 2017. This study shows that the community composition of viable diatom resting stages is largely affected by the timing of the sea ice retreat. [ABSTRACT FROM AUTHOR]

Satellite‐derived data suggest an increase in annual primary production following the loss of summer sea ice in the Arctic Ocean. The scarcity of field data to corroborate this enhanced algal production incited a collaborative project combining six annual cycles of sequential sediment trap measurements obtained over a 17‐year period in the Eurasian Arctic Ocean. Here we present microalgal fluxes measured at ~200 m to reflect the bulk of algal carbon production. Ice algae contributed to a large proportion of the microalgal carbon export before complete ice melt and possible detection of their production by satellites. In the northern Laptev Sea, annual microalgal carbon fluxes were lower during the 2007 minimum ice extent than in 2006. In 2012, early snowmelt led to early microalgal carbon flux in the Nansen Basin. Hence, a change in the timing of snowmelt and ice algae release may affect productivity and export over the Arctic basins. Plain Language Summary: An increase in algal production has been reported following the recent decline in sea ice cover in the Arctic Ocean. However, very few field measurements have been conducted to corroborate these remote sensing‐derived observations. Algal cell fluxes collected with sediment traps at ~200 m during six annual cycles between 1995 and 2012 were combined to determine changes in the timing, magnitude, and composition of the algal export in the Eurasian Arctic Ocean. Algal fluxes, mostly composed of ice algae, reflected a large fraction of the surface phytoplankton carbon production. Ice algae were nearly all collected during snowmelt and before complete ice melt, preventing the detection of their production by satellites. While relatively high algal export was observed under thick ice cover in 1996, early algal export in 2012 suggested an early release of ice algae due to early snowmelt. Overall, snow cover played an important role in regulating the timing and magnitude of ice algae release in the Eurasian Arctic Ocean. Key Points: The majority of algal cells were exported before complete ice meltEarly snowmelt led to the early release of ice algae prior to the record minimum sea ice extent of 2012Phytoplankton carbon contributed up to ~40% of the particulate organic carbon fluxes during short intense export events [ABSTRACT FROM AUTHOR]

During ice melt in spring, ice algae are released from the ice and could be exposed to variable temperatures and irradiances in surface water. Saroma Lagoon is an embayment with two inlets leading to the Sea of Okhotsk. With seasonal development of sea ice, its water temperature changes dramatically throughout the year. To investigate the living and photoprotective strategies of ice algae in such a coastal water system, we grew Nitzschia cf. neglecta, an ice diatom isolated from the sea ice of this lagoon, under irradiance levels of 30 and 100 lmol photons • m-2 • s-1, and temperatures of 2°C and 10°C. Then the acclimated cells were exposed to high light in order to investigate the plasticity of their photosynthetic apparatus. At 10°C, cells grew faster and showed decreased susceptibility to high light. At 2°C, an immediate decrease in all pigment content upon exposure, as well as a higher cellular content of diatoxanthin was used to compensate for the more severe excitation stress. Highly efficient photoprotection was achieved through the diadinoxanthin-diatoxanthin cycledependent nonphotochemical quenching. While regulation through psbA and rbcL at the transcription level played a minor role in the response to high light stress at both temperatures. The wide tolerance to both temperature and light changes suggest that the thinning of sea ice and higher temperatures in a warmer world will lead to more intense blooms in Saroma Lagoon. [ABSTRACT FROM AUTHOR]

A: In a match situation, ice algae production and release precede the pelagic phytoplankton bloom. Capital breeder C. hyperboreus reproduce at depth before the spring ascent. Buoyant eggs develop into nauplii during their ascent, reaching surface waters in time for the bloom. Income breeder C. glacialis match ice algae production to fuel their maturation and egg production while nauplii feed on the ensuing pelagic bloom to fuel their growth. Older copepodites start their downward migration for overwintering at the end of the productive season and before ice formation. B: In a mismatch situation such as the one caused by the record minimum sea ice extent of 2012, the extended open water season allowed enhanced solar absorption. Warmer winter water temperature allowed C. hyperboreus females to overwinter and spawn at shallower depths than usual, resulting in a peak in nauplii abundance 12–14 weeks before the diatom bloom. The late sea ice break-up and snow melt due to the ice rebound of 2013 delayed the ice algae bloom and export, causing a 13–15-week mismatch with C. glacialis upward migration. The overlapping export of ice and pelagic diatoms suppressed the gap usually exploited by C. glacialis for the development of their nauplii into the first feeding stage at the time of pelagic production. These events led to the lowest summertime nauplii abundance recorded, implying a poor copepod recruitment the following year. • Record sea ice minimum in 2012 led to warmer water temperature in the Beaufort Sea. • Anomalous sea ice growth in 2013 delayed ice algae release and pelagic production. • These anomalies disrupted the timing between microalgae and herbivorous copepods. The unprecedented pace of warming of the Arctic Ocean affects a wide range of pelagic processes, from primary production to fish recruitment. Sequential sediment traps deployed on oceanographic moorings at the Beaufort Sea shelf break were used to investigate the impact of variations in physical conditions on the phenology of microalgae and herbivorous copepods. Water temperature, salinity, microalgal fluxes, and zooplankton fluxes were monitored over 6 of the 7 annual cycles from September 2009 to September 2016 with a hiatus from September 2013 to August 2014. Satellite-derived sea ice concentration and snow depth were retrieved for the same period. For 5 of the 6 cycles, a consistent interval between the vernal abundance of the Calanus species (C. glacialis and C. hyperboreus), the export of the ice alga Nitzschia frigida, and peak in nauplii abundance and diatom export, reflected a match between microalgae and herbivorous copepods. Drastic changes in water temperatures and ice conditions during the 2012–2013 cycle affected this synchronization. In 2012, the record sea ice minimum extent led to anomalously high upper water temperatures during autumn and winter that led to a shoaling of the vertical distribution of C. hyperboreus females and to the production of their nauplii at shallower depths several weeks before diatom export. Due to several oceanic and atmospheric forcing, the record sea ice minimum extent was followed in 2013 by a rebound in ice cover that delayed snowmelt, sea ice melt, and ice algae export. This delay resulted in a mismatch between C. glacialis females and N. frigida fluxes and in fewer C. glacialis nauplii in June and July 2013. These mismatch events likely had a negative impact on Calanus recruitment and on the subsequent transfer of energy to carnivorous zooplankton and fish. It is crucial to maintain mooring deployments to accurately assess the wide range of effects of climate warming on the marine ecosystems. [ABSTRACT FROM AUTHOR]

Recent studies have highlighted the importance of ice-algal blooms in driving darkening and therefore surface melt of the Greenland Ice Sheet (GrIS). However, the contribution of fungal and bacterial communities to this microbially driven albedo reduction remains unconstrained. To address this significant knowledge gap, fungi were isolated from key GrIS surface habitats (surface ice containing varying abundance of ice algae, supraglacial water, cryoconite holes, and snow), and a combination of cultivation and sequencing methods utilized to characterize the algal-associated fungal and bacterial diversity and abundance. Six hundred and ninety-seven taxa of fungi were obtained by amplicon sequencing and more than 200 fungal cultures belonging to 46 different species were isolated through cultivation approaches. Basidiomycota dominated in surface ice and water samples, and Ascomycota in snow samples. Amplicon sequencing revealed that bacteria were characterized by a higher diversity (883 taxa detected). Results from cultivation as well as ergosterol analyses suggested that surface ice dominated by ice algae and cryoconite holes supported the highest fungal biomass (104–105 CFU/100 ml) and that many fungal taxa recognized as endophytes and plant pathogens were associated with dark ice characterized by a high abundance of ice algae. This paper significantly advances this field of research by investigating for the first time the fungal abundance and diversity associated with algal blooms causing the darkening of the GrIS. There is a strong association between the abundance and diversity of fungal species and the blooming of algae on the surface ice of the Greenland Ice Sheet. [ABSTRACT FROM AUTHOR]

During the summer ablation season, Arctic glacier surfaces host a wealth of microbial life. Here, the phototroph communities on the ablating bare-ice surface of three valley glaciers on Brøggerhalvøya, Svalbard were investigated. The communities mainly comprised seven taxa of green algae and cyanobacteria, which have been commonly reported on Arctic glaciers. Although the geographical and glaciological settings of the three studied glaciers are similar, there were differences in total phototroph biomass. The community structure was also distinctive among the glaciers: high dominance of a single taxon of green algae (Ancylonema nordenskiöldii) for Midtre Lovénbreen, abundant cyanobacteria for Austre Brøggerbreen, and diverse green algae for Pedersenbreen. The major soluble ions in the surface ice showed that there was no significant difference in meltwater nutrient conditions between the glaciers, but there were lower concentrations of mineral-derived ions on Midtre Lovénbreen. Consequently, the glacier-specific mineral loading and surface hydrology are inferred to explain the contrast in bare ice algal communities between the glaciers. We hypothesize that local, glacier-specific conditions affect algal communities and the associated influences on carbon cycling and ice-surface albedo.

During the N-ICE2015 drift expedition north-west of Svalbard, we observed the establishment and development of algal communities in first-year ice (FYI) ridges and at the snow-ice interface. Despite some indications of being hot spots for biological activity, ridges are under-studied largely because they are complex structures that are difficult to sample. Snow infiltration communities can grow at the snow-ice interface when flooded. They have been commonly observed in the Antarctic, but rarely in the Arctic, where flooding is less common mainly due to a lower snow-to-ice thickness ratio. Combining biomass measurements and algal community analysis with under-ice irradiance and current measurements as well as light modeling, we comprehensively describe these two algal habitats in an Arctic pack ice environment. High biomass accumulation in ridges was facilitated by complex surfaces for algal deposition and attachment, increased light availability, and protection against strong under-ice currents. Notably, specific locations within the ridges were found to host distinct ice algal communities. The pennate diatoms Nitzschia frigida and Navicula species dominated the underside and inclined walls of submerged ice blocks, while the centric diatom Shionodiscus bioculatus dominated the top surfaces of the submerged ice blocks. Higher light levels than those in and below the sea ice, low mesozooplankton grazing, and physical concentration likely contributed to the high algal biomass at the snow-ice interface. These snow infiltration communities were dominated by Phaeocystis pouchetii and chain-forming pelagic diatoms (Fragilariopsis oceanica and Chaetoceros gelidus). Ridges are likely to form more frequently in a thinner and more dynamic ice pack, while the predicted increase in Arctic precipitation in some regions in combination with the thinning Arctic icescape might lead to larger areas of sea ice with negative freeboard and subsequent flooding during the melt season. Therefore, these two habitats are likely to become increasingly important in the new Arctic with implications for carbon export and transfer in the ice-associated ecosystem.

Krill overwintering strategies vary with ontogeny and year; understanding this variability is essential to predicting how the species will respond to climate change in the future. Overwintering studies have focused on larval and adult krill, but we know little about how juvenile krill overwinter. The late winter diet of juvenile krill is important because it will determine their growth and development rates and consequently their reproductive potential the following spring. A diet rich in ice algae would promote growth and reproductive development. The Bransfield Strait (northern Antarctic Peninsula, AP) is an important overwintering ground for krill; it has been proposed this region offers a food-rich winter environment. We examined the contribution of ice algae to the energy budget of overwintering juvenile krill during 2 years with contrasting sea ice conditions. Grazing on ice algae contributed ∼146% to their winter energy budget in 2015, even though ice concentrations were ≤50% and consisted of newly formed pancake ice. However, when sea ice advanced late in the Bransfield Strait (2016), ice algae contributed significantly less (∼16%) to the winter energy budget of juvenile krill. Delayed sea ice advance may negatively affect growth and reproductive development of overwintering juvenile krill. [ABSTRACT FROM AUTHOR]

Historical sea ice core chlorophyll‐a (Chla) data are used to describe the seasonal, regional, and vertical distribution of ice algal biomass in Antarctic landfast sea ice. The analyses are based on the Antarctic Fast Ice Algae Chlorophyll‐a data set, a compilation of currently available sea ice Chla data from landfast sea ice cores collected at circum‐Antarctic nearshore locations between 1970 and 2015. Ice cores were typically sampled from thermodynamically grown first‐year ice and have thin snow depths (mean = 0.052 ± 0.097 m). The data set comprises 888 ice cores, including 404 full vertical profile cores. Integrated ice algal Chla biomass (range: <0.1–219.9 mg/m2, median = 4.4 mg/m2, interquartile range = 9.9 mg/m2) peaks in late spring and shows elevated levels in autumn. The seasonal Chla development is consistent with the current understanding of physical drivers of ice algal biomass, including the seasonal cycle of irradiance and surface temperatures driving landfast sea ice growth and melt. Landfast ice regions with reported platelet ice formation show maximum ice algal biomass. Ice algal communities in the lowermost third of the ice cores dominate integrated Chla concentrations during most of the year, but internal and surface communities are important, particularly in winter. Through comparison of biomass estimates based on different sea ice sampling strategies, that is, analysis of full cores versus bottom‐ice section sampling, we identify biases in common sampling approaches and provide recommendations for future survey programs: for example, the need to sample fast ice over its entire thickness and to measure auxiliary physicochemical parameters. Plain Language Summary: Antarctic sea ice is a key driver of physical, chemical, and biological processes in the Southern Ocean. Importantly, sea ice serves as a substrate for microscopic algae which grow in the bottom, interior, and surface layers of the ice. These algae are considered an important food source for Antarctic marine food webs. Using a newly collated database of historical sea ice core chlorophyll‐a data (a proxy for ice algal biomass) from coastal sites, we describe the seasonal and vertical variability of algal biomass in Antarctic landfast sea ice. The seasonal chlorophyll‐a development is consistent with the current understanding of physical drivers of ice algal biomass, including the seasonal cycle of irradiance and surface temperatures driving landfast sea ice growth and melt. Our analyses show that algae in the lowermost third of ice cores drive the annual cycle of integrated biomass, but internal and surface communities are also important. Through comparison of biomass estimates based on different sea ice sampling strategies, that is, analysis of full cores versus bottom‐ice section sampling, we identify biases in common sampling approaches and provide recommendations for future survey programs: for example, the need to sample fast ice over its entire thickness and to measure auxiliary physical parameters, in particular snow‐thickness data. Key Points: First comprehensive collation of Antarctic landfast sea‐ice core chlorophyll‐a data establishedAlgae in lowermost third of ice cores drive annual cycle of integrated biomass, but internal and surface communities are also importantFull‐profile ice‐core sampling and systematic collection of auxiliary physico‐chemical parameters is recommended for future studies [ABSTRACT FROM AUTHOR]

Sea ice provides an important habitat for the ice algae that fuel upper trophic levels in early spring, prior to phytoplankton bloom development. In the Chukchi Sea, the ice environment has experienced large‐scale changes in snow cover, ice thickness and extent, and timing of ice advance and retreat. Using a 1‐D coupled physical‐biological ice‐ecosystem model and observed distributions of sea ice, we investigated how changing ice conditions may have impacted ice algal bloom dynamics for the central Chukchi Sea between 1980 and 2015. Model results suggest that over this 35‐year time period, annual ice algal net primary production (NPP) decreased by 22%. Modeled snow and ice melt, as well as satellite‐derived ice retreat, occurred progressively earlier in the spring from 1980 to 2015, even though modeled ice and snow thicknesses showed no temporal trends. Over time, ice algal blooms ended earlier in spring due to earlier onset of ice melt and ice retreat. Results suggest that the length of the ice season sets the upper bound on annual ice algal NPP, with late melt years corresponding to highest NPP. Linear regression results suggest that within early versus late melt year groupings, snow plays a secondary, yet important role in determining annual ice algal NPP. As sea ice thins and melts earlier in spring, future snow dynamics will be an important factor determining the onset of the ice algal bloom and thus the duration of the bloom in an early melt, shortened ice season scenario. Plain Language Summary: Sea ice provides an important habitat for the ice algae that fuel upper trophic levels in early spring, prior to phytoplankton bloom development. In the Chukchi Sea, the ice environment has experienced large‐scale changes in snow cover, ice thickness and extent, and timing of ice advance and retreat. Using a modeling study that simulated the sea ice environment and sea ice ecosystem, we investigated how changing ice conditions may have impacted ice algal bloom dynamics for the central Chukchi Sea between 1980 and 2015. Modeling results suggest that ice algal blooms end earlier in spring due to earlier onset of ice melt and ice retreat. Earlier bloom termination leads to declines in ice algal production between 1980 and 2015. Key Points: Ice algal production has decreased 22% in the central Chukchi Sea from 1980 to 2015Ice season duration sets the upper limit on annual ice algal productionSnow thickness can compensate for or amplify the impact of a shortening ice season [ABSTRACT FROM AUTHOR]

It is fundamental to understand the development of Zygnematophycean (Streptophyte) micro-algal blooms within Greenland Ice Sheet (GrIS) supraglacial environments, given their potential to significantly impact both physical (melt) and chemical (carbon and nutrient cycling) surface characteristics. Here, we report on a space-for-time assessment of a GrIS ice algal bloom, achieved by sampling an ~85 km transect spanning the south-western GrIS bare ice zone during the 2016 ablation season. Cell abundances ranged from 0 to 1.6 x 104 cells ml-1, with algal biomass demonstrated to increase in surface ice with time since snow line retreat (R² = 0.73, P < 0.05). A suite of light harvesting and photo-protective pigments were quantified across transects (chlorophylls, carotenoids and phenols) and shown to increase in concert with algal biomass. Ice algal communities drove net autotrophy of surface ice, with maximal rates of net production averaging 0.52 ± 0.04 mg C 1-1 d-1, and a total accumulation of 1.306 Gg C (15.82 ± 8.14 kg C km-2) predicted for the 2016 ablation season across an 8.24 x 104 km² region of the GrIS. By advancing our understanding of ice algal bloom development, this study marks an important step toward projecting bloom occurrence and impacts into the future. [ABSTRACT FROM AUTHOR]

Red-colored secondary pigments in glacier algae play an adaptive role in melting snow and ice. We advance this hypothesis using a model of color-based absorption of irradiance, an experiment with colored particles in snow, and the natural history of glacier algae. Carotenoids and phenols--astaxanthin in snow-algae and purpurogallin in ice-algae--shield photosynthetic apparatus by absorbing overabundant visible wavelengths, then dissipating the excess radiant energy as heat. This heat melts proximal ice crystals, providing liquid-water in a 0°C environment and freeing up nutrients bound in frozen water. We show that purple-colored particles transfer 87%-89% of solar energy absorbed by black particles. However, red-colored particles transfer nearly as much (85%-87%) by absorbing peak solar wavelengths and reflecting the visible wavelengths most absorbed by nearby ice and snow crystals; this latter process may reduce potential cellular overheating when snow insulates cells. Blue and green particles transfer only 80%-82% of black particle absorption. In the experiment, red-colored particles melted 87% as much snow as black particles, while blue particles melted 77%. Green-colored snow-algae naturally occupy saturated snow where water is non-limiting; red-colored snow-algae occupy drier, water-limited snow. In addition to increasing melt, we suggest that esterified astaxanthin in snow-alga cells increases hydrophobicity to remain surficial. [ABSTRACT FROM AUTHOR]

Abstract: Fast ice is an important component of Antarctic coastal marine ecosystems, providing a prolific habitat for ice algal communities. This work examines the relationships between normalized difference indices (NDI) calculated from under‐ice radiance measurements and sea ice algal biomass and snow thickness for Antarctic fast ice. While this technique has been calibrated to assess biomass in Arctic fast ice and pack ice, as well as Antarctic pack ice, relationships are currently lacking for Antarctic fast ice characterized by bottom ice algae communities with high algal biomass. We analyze measurements along transects at two contrasting Antarctic fast ice sites in terms of platelet ice presence: near and distant from an ice shelf, i.e., in McMurdo Sound and off Davis Station, respectively. Snow and ice thickness, and ice salinity and temperature measurements support our paired in situ optical and biological measurements. Analyses show that NDI wavelength pairs near the first chlorophyll a (chl a) absorption peak (≈440 nm) explain up to 70% of the total variability in algal biomass. Eighty‐eight percent of snow thickness variability is explained using an NDI with a wavelength pair of 648 and 567 nm. Accounting for pigment packaging effects by including the ratio of chl a‐specific absorption coefficients improved the NDI‐based algal biomass estimation only slightly. Our new observation‐based algorithms can be used to estimate Antarctic fast ice algal biomass and snow thickness noninvasively, for example, by using moored sensors (time series) or mapping their spatial distributions using underwater vehicles. [ABSTRACT FROM AUTHOR]

Assessing the role of sea ice algal biomass and primary production for polar ecosystems remains challenging due to the strong spatio-temporal variability of sea ice algae. Therefore, the spatial representativeness of sea ice algal biomass and primary production sampling remains a key issue in large-scale models and climate change predictions of polar ecosystems. To address this issue, we presented two novel approaches to up-scale ice algal chl a biomass and net primary production (NPP) estimates based on profiles covering distances of 100 to 1,000 s of meters. This was accomplished by combining ice core-based methods with horizontal under-ice spectral radiation profiling conducted in the central Arctic Ocean during summer 2012. We conducted a multi-scale comparison of ice-core based ice algal chl a biomass with two profiling platforms: a remotely operated vehicle and surface and under ice trawl (SUIT). NPP estimates were compared between ice cores and remotely operated vehicle surveys. Our results showed that ice core-based estimates of ice algal chl a biomass and NPP do not representatively capture the spatial variability compared to the remotely operated vehicle-based estimates, implying considerable uncertainties for pan-Arctic estimates based on ice core observations alone. Grouping sea ice cores based on region or ice type improved the representativeness. With only a small sample size, however, a high risk of obtaining non-representative estimates remains. Sea ice algal chl a biomass estimates based on the dominant ice class alone showed a better agreement between ice core and remotely operated vehicle estimates. Grouping ice core measurements yielded no improvement in NPP estimates, highlighting the importance of accounting for the spatial variability of both the chl a biomass and bottom-ice light in order to get representative estimates. Profile-based measurements of ice algae chl a biomass identified sea ice ridges as an underappreciated component of the Arctic ecosystem because chl a biomass was significantly greater in this unique habitat. Sea ice ridges are not easily captured with ice coring methods and thus require more attention in future studies. Based on our results, we provide recommendations for designing an efficient and effective sea ice algal sampling program for the summer season.

Antarctic krill Euphausia superba (“krill”) constitute a fundamental food source for Antarctic seabirds and mammals, and a globally important fisheries resource. The future resilience of krill to climate change depends critically on the winter survival of young krill. To survive periods of extremely low production by pelagic algae during winter, krill are assumed to rely partly on carbon produced by ice algae. The true dependency on ice algae-produced carbon, however, is so far unquantified. This confounds predictions on the future resilience of krill stocks to sea ice decline. Fatty acid (FA) analysis, bulk stable isotope analysis (BSIA), and compound-specific stable isotope analysis (CSIA) of diatom- and dinoflagellate-associated marker FAs were applied to quantify the dependency of overwintering larval, juvenile, and adult krill on ice algae-produced carbon (αIce) during winter 2013 in the Weddell-Scotia Confluence Zone. Our results demonstrate that the majority of the carbon uptake of the overwintering larval and juvenile krill originated from ice algae (up to 88% of the carbon budget), and that the dependency on ice algal carbon decreased with ontogeny, reaching

Abstract: Microalgae colonizing the underside of sea ice in spring are a key component of the Arctic foodweb as they drive early primary production and transport of carbon from the atmosphere to the ocean interior. Onset of the spring bloom of ice algae is typically limited by the availability of light, and the current consensus is that a few tens‐of‐centimeters of snow is enough to prevent sufficient solar radiation to reach underneath the sea ice. We challenge this consensus, and investigated the onset and the light requirement of an ice algae spring bloom, and the importance of snow optical properties for light penetration. Colonization by ice algae began in May under >1 m of first‐year sea ice with ∼1 m thick snow cover on top, in NE Greenland. The initial growth of ice algae began at extremely low irradiance (<0.17 μmol photons m−2 s−1) and was documented as an increase in Chlorophyll a concentration, an increase in algal cell number, and a viable photosynthetic activity. Snow thickness changed little during May (from 110 to 91 cm), however the snow temperature increased steadily, as observed from automated high‐frequency temperature profiles. We propose that changes in snow optical properties, caused by temperature‐driven snow metamorphosis, was the primary driver for allowing sufficient light to penetrate through the thick snow and initiate algae growth below the sea ice. This was supported by radiative‐transfer modeling of light attenuation. Implications are an earlier productivity by ice algae in Arctic sea ice than recognized previously. [ABSTRACT FROM AUTHOR]

A numerical mechanism connecting ice algal ecodynamics with the buildup of organic macromolecules is tested within modeled pan-Arctic brine channels. The simulations take place offline in a reduced representation of sea ice geochemistry. Physical driver quantities derive from the global sea ice code CICE, including snow cover, thickness and internal temperature. The framework is averaged over ten boreal biogeographic zones. Computed nutrient-light-salt limited algal growth supports grazing, mortality and carbon flow. Vertical transport is diffusive but responds to pore structure. Simulated bottom layer chlorophyll maxima are reasonable, though delayed by about a month relative to observations due to uncertainties in snow variability. Upper level biota arise intermittently during flooding events. Macromolecular concentrations are tracked as proxy proteins, polysaccharides, lipids and refractory humics. The fresh biopolymers undergo succession and removal by bacteria. Baseline organics enter solely through cell disruption, thus the internal carbon content is initially biased low. By including exudation, agreement with dissolved organic or individual biopolymer data is achieved given strong release coupled to light intensity. Detrital carbon then reaches hundreds of micromolar, sufficient to support structural changes to the ice matrix. [ABSTRACT FROM AUTHOR]

We determined, and compared, the abiotic stress state of algae and their attached bacterial communities following their release into the water column during the Arctic sea ice melt season using specific lipid markers that are characteristic of type II photo-oxidation processes, together with those associated with cis-trans isomerase and 10 S -DOX-like lipoxygenase activity, which are indicative of salinity stress. More specifically, parent lipids and some of their oxidation products were quantified in sinking particles (from sediment traps) collected from the Beaufort Sea, Resolute Passage and the Davis Strait (Canadian Arctic) in 2009, 2012 and 2015, respectively. Our data show that salinity- and light-induced bacterial stress processes are temporally decoupled, with the former occurring at the beginning of ice melting and the latter observed during the final stages of ice melt and in subsequent open waters. The salinity-induced bacterial stress seen during the early stages of ice melting contrasts the healthy state of sea ice algae under the same hypersaline conditions. As sea ice melt progresses, brine channels become hyposaline, thus reducing the bacterial salinity stress, while ice algae become susceptible to photodegradation by singlet oxygen ( 1 O 2 ), despite relatively low temperatures and irradiance. However, the presence of high amounts of extracellular polymeric substances (EPS) encapsulating the algae likely prevents the diffusion of 1 O 2 to the attached bacteria. Photo-oxidative stress is enhanced further in open waters due to the low amounts of EPS. The lower amounts of active bacteria in the Arctic compared to temperate or tropical regions is thus attributed to the combined action of efficient salinity stress in spring and photo-oxidative stress in summer. Outcomes from this study also imply that surface sediments from the Canadian Arctic contain relatively high quantities of sea ice-derived organic matter that has undergone a strong degree of salinity-induced stress. [ABSTRACT FROM AUTHOR]