Search

We studied phytoplankton and protozooplankton community composition based on light microscopy, flow cytometry, and photosynthetic pigment data in the Atlantic sector of the Southern Ocean during March 2019 (early austral autumn). Sampling was focused on the area east of the prime meridian in the Kong Håkon VII Hav, including Astrid Ridge, Maud Rise, and a south–north transect at 6∘ E. Phytoplankton community composition throughout the studied area was characterized by oceanic diatoms typical of the iron-depleted high-nutrient, low-chlorophyll (HNLC) Southern Ocean. Topography and wind-driven iron supply likely sustained blooms dominated by the centric diatom Chaetoceros dichaeta at Maud Rise and at a station north of the 6∘ E transect. For the remainder of the 6∘ E transect, diatom composition was similar to the previously mentioned bloom stations, but flagellates dominated in abundance, suggesting a post-bloom situation and likely top-down control by krill on the bloom-forming diatoms. Among flagellates, species with haptophyte-type pigments were the dominating group. At Astrid Ridge, overall abundances were lower and pennate diatoms were more numerous than centric diatoms, but the community composition was nevertheless typical of HNLC areas. The observations described here show that C. dichaeta can form blooms beyond the background biomass level and also fuels both carbon export and upper trophic levels within HNLC areas. This study is the first thorough assessment of phytoplankton communities in this region and can be compared to other seasons in future studies.

Seasonal plankton time-series data are presented from Kongsfjorden from two years with contrasting environmental conditions. Kongsfjorden (west coast of Spitsbergen – 79◦N) integrates inputs from Atlantic and Arctic waters, and glacier run-off, and is thus a prime location to study impacts on ecosystem dynamics of key environmental drivers that are relevant across the Arctic. Despite extensive research in Kongsfjorden, seasonally resolved data are scarce. From late April/early May to early September 2019 and 2020, we conducted pelagic sampling at a mid-fjord station at mostly weekly to bi-weekly resolution investigating the environmental drivers of phyto- and zooplankton community composition and phenology. During spring 2019, Atlantic water masses with temperatures > 1 ◦C were found throughout the upper 250 m of the water column, and little sea ice occurred in the fjord. Spring 2020, in turn, was characterized by the presence of local water masses with sub-zero temperatures and relatively extensive sea-ice cover. The most striking contrast between the two years was the difference in phytoplankton spring bloom composition. In 2019, the spring bloom was dominated by the colonial stage of the haptophyte Phaeocystis pouchetii and diatoms played a minor role, while the spring bloom in 2020 was dominated by diatoms of the genus Thalassiosira succeeded by P. pouchetii. Selective grazing by large copepods and water mass structure seem to have been the decisive factors explaining the marked difference in diatom spring bloom biomass between the years while similar spring abundances of P. pouchetii in both years indicated that this species was less impacted by those factors. Our data suggest that differences in spring bloom composition impacted trophic transfer and carbon export. Recruitment of the dominant copepods Calanus finmarchicus and C. glacialis, Cirripedia and euphausiid larvae as well as the export of carbon to the seabed was more efficient during the diatom-dominated compared to the P. pouchetii–dominated spring bloom. In summer, the plankton composition shifted towards a flagellate-dominated community characterized by mixo- and heterotrophic taxa adapted to a lower nutrient regime and strong top-down control by copepod grazers. However, residual silicic acid after the P. pouchetii–dominated spring bloom fueled a late summer diatom bloom in 2019. Our data provide a first glimpse into the environmental drivers of plankton phenology and underline that high resolution monitoring over many annual cycles is required to resolve the ephemeral variations of plankton populations against the backdrop of climate change.

Ongoing climate change in the Arctic has caused tidewater glaciers to retreat while increasing the discharge of freshwater and terrestrial material into fjords. This can affect both nutrient inputs and cycling within the fjord systems. In particular, tidewater glaciers and the presence of associated subglacial meltwater plumes can have a large impact on fjord circulation and biogeochemistry. In this study, we assess the influence of tidewater glaciers on nitrogen inputs and cycling in two fjords in Svalbard during the summer using stable isotopic analyses of dissolved nitrate (δ15N and δ18O) in combination with nutrient and hydrographic data. Kongsfjorden receives inputs from tidewater glaciers, whereas Rijpfjorden mainly receives surface inputs from land-terminating glaciers. Results showed that both fjords are enriched in nutrients from terrestrial inputs. Nutrient ratios indicate excess Si and P relative to N. In both fjords, terrestrial nitrate from snowpack and glacier melting are identified as the dominant sources based on high δ18O-NO3- and low δ15N-NO3- of dissolved nitrate. In Kongsfjorden, mixed-layer nitrate is completely consumed within the fjord system, which we attribute to vigorous circulation at the glacial front influenced by the subglacial plume and longer residence time in the fjord. This is in contrast to Rijpfjorden where nutrients are only partially consumed perhaps due to surface river discharge and light limitation. In Kongsfjorden, we estimate terrestrial and marine N contributions to the nitrate pool from nitrogen isotopic values (δ15N-NO3-), and this suggests that nearly half the nitrate in the subglacial plume (50 ± 3 %) and the water column (44 ± 3 %) originates from terrestrial sources. We show that terrestrial N contributes significantly to the regenerated N pool (63 %–88 %) within this fjord suggesting its importance in sustaining productivity here. Given this importance of terrestrial nutrient sources within the fjords, increase in these inputs due to climate change can enhance the fjord nutrient inventory, productivity and nutrient export offshore. Specifically, increasing Atlantification and warmer Atlantic Water will encourage tidewater glacier retreat and in turn increase surface discharge. In fjords akin to Rijpfjorden this is expected to foster more light limitation and less dynamic circulation, ultimately aiding the export of nutrients offshore contributing to coastal productivity. Climate change scenarios postulated for fjords such as Kongsfjorden include more terrestrial N-fuelled productivity and N cycling within the fjord, less vigorous circulation due to the retreat of tidewater glaciers, and the expansion of oxygen-depleted deep waters isolated by the sill.

Microalgae growing within and attached to the bottom of Arctic sea ice (sympagic algae) can serve as a nutritious food resource for animals inhabiting the sea-ice water interface (under-ice fauna), particularly during the bottom ice-algal bloom in spring. As a consequence, under-ice fauna is likely impacted by sea-ice decline and changes in ice-algal primary production. To investigate this, samples of pelagic (=PPOM) and ice-associated particulate organic matter (=IPOM) and the ice-associated amphipods Apherusa glacialis and Eusirus holmii, and polar cod (Boreogadus saida), collected below ridged sea ice at two locations with pronounced differences in productivity in the northern Barents Sea during May 2021, were assessed for their trophic marker content. Specifically, we investigated the composition of diatom- and dinoflagellate-produced fatty acids (FAs), pelagic and sympagic highly branched isoprenoid (HBI) lipids as well as sterols to determine the animals’ dietary preferences and trophic association to the sea-ice habitat during spring. Relative proportions of FAs differed strongly between PPOM and IPOM, indicating differences in species composition and degradation state between pelagic and sympagic habitats, respectively. FA signatures and sterol content of the consumers largely resembled known diet compositions with a strong reliance on diatom-derived carbon in A. glacialis, a higher degree of carnivory in E. holmii and evidence of Calanus-feeding in polar cod. Sympagic HBIs were detected at either low concentrations or not at all, in both producers and consumers, likely as a result of the very low abundance of their source diatoms. Pronounced trophic marker variability in A. glacialis collected at the highly productive shelf slope station versus the less productive central Arctic Basin station suggests a surprisingly high flexibility in carbon-source composition with a stronger reliance on pelagic food when available versus a higher importance of ice algal carbon when pelagic production is low. Nevertheless and despite the general lack (below detection limit) of sympagic HBIs in our dataset, high ice-algal biomass and elevated proportions of polyunsaturated FAs in IPOM compared to other seasons indicate that ice algae constitute a valuable nutritional carbon source as alternative to pelagic carbon during spring.

The Arctic sea-ice-scape is rapidly transforming. Increasing light penetration will initiate earlier seasonal primary production. This earlier growing season may be accompanied by an increase in ice algae and phytoplankton biomass, augmenting the emission of dimethylsulfide and capture of carbon dioxide. Secondary production may also increase on the shelves, although the loss of sea ice exacerbates the demise of sea-ice fauna, endemic fish and megafauna. Sea-ice loss may also deliver more methane to the atmosphere, but warmer ice may release fewer halogens, resulting in fewer ozone depletion events. The net changes in carbon drawdown are still highly uncertain. Despite large uncertainties in these assessments, we expect disruptive changes that warrant intensified long-term observations and modelling efforts., info:eu-repo/semantics/published

Despite the exclusion of the Southern Ocean from assessments of progress towards achieving the Convention on Biological Diversity (CBD) Strategic Plan, the Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR) has taken on the mantle of progressing efforts to achieve it. Within the CBD, Aichi Target 11 represents an agreed commitment to protect 10% of the global coastal and marine environment. Adopting an ethos of presenting the best available scientific evidence to support policy makers, CCAMLR has progressed this by designating two Marine Protected Areas in the Southern Ocean, with three others under consideration. The region of Antarctica known as Dronning Maud Land (DML; 20°W to 40°E) and the Atlantic sector of the Southern Ocean that abuts it conveniently spans one region under consideration for spatial protection. To facilitate both an open and transparent process to provide the vest available scientific evidence for policy makers to formulate management options, we review the body of physical, geochemical and biological knowledge of the marine environment of this region. The level of scientific knowledge throughout the seascape abutting DML is polarized, with a clear lack of data in its eastern part which is presumably related to differing levels of research effort dedicated by national Antarctic programmes in the region. The lack of basic data on fundamental aspects of the physical, geological and biological nature of eastern DML make predictions of future trends difficult to impossible, with implications for the provision of management advice including spatial management. Finally, by highlighting key knowledge gaps across the scientific disciplines our review also serves to provide guidance to future research across this important region.

Recent reports on Arctic underice phytoplankton blooms have directed attention to primary production below the sea ice cover. Such underice blooms cannot be detected from space; thus, methods for autonomous underice measurements are critically needed to extend observations beyond ship-based surveys. One central aspect of the ecology of these blooms is whether they were advected from open-water areas or were able to develop below the ice cover under typically low light conditions. The photoacclimation state of the bloom can provide clues about the growth conditions and therefore its origin. Here we investigate the photoacclimation state of a Phaeocystis pouchetii-dominated underice bloom in the Arctic Ocean using ratios of photoprotective carotenoids (PPC) to photosynthetic carotenoids (PSC) and chlorophyll a. The pigment proxies indicate local growth under the ice pack. Furthermore, a method using in situ light absorption measurements to estimate the PPC:PSC ratio was in agreement with the pigment data. The slope of in situ phytoplankton absorption between 488 and 532nm, affected by both PPC and PSC, had a significant linear relationship to the PPC:PSC ratio, indicating that prediction of photoacclimation state can be obtained from absorption profiles. We also review, with regard to the pigment function, different ways of grouping pigments into PPC or PSC applied in previous studies. Although more validation data sets are needed to assess the impact of pigment packaging on the relationship between PPC:PSC and absorption measurement slopes, our study shows the potential for using in situ absorption measurements to collect information about phytoplankton physiology below sea ice. Plain Language Summary Phytoplankton blooms below sea ice cover, that is, underice blooms, can be advected by ocean currents below the ice pack from ice-free waters where they had sufficient light available for growth. They can also grow below the ice pack if sufficient light is transmitted through the ice, and they are able to adjust to the typically low light conditions. Algal pigments can indicate which of the two scenarios is more likely. Algal pigments capture sunlight for photosynthesis, but some of them are used to protect the algal cells from excess light. The ratio of photoprotective to photosynthetic pigments reveals the photoacclimation state ("short time adjustments") of the algae. In this study we investigate the photoacclimation state of an Arctic underice bloom based on pigment ratios, which indicate local growth below the ice pack. Furthermore, we validate a method to estimate the pigment ratios from light absorption measurements carried out with an instrument in the water column. Studies like these are important for developing methods to study remote areas. Ship-based surveys can only cover a restricted area, and satellite remote sensing does not provide biological information from ice-covered waters. Our study shows promising results for using water column instruments to study phytoplankton physiology.

The Arctic Ocean is facing rapid environmental changes with cascading effects on the entire Arctic marine ecosystem. However, we have a limited understanding of the consequences such changes have on bacteria and archaea (prokaryotes) at the base of the marine food web. In this study, we show how the prokaryotic rare biosphere behaves over a range of highly heterogeneous environmental conditions using 16S rRNA gene reads from amplicon and metagenome sequencing data from seawater samples collected during the Norwegian young sea ICE expedition between late winter and early summer. The prokaryotic rare biosphere was analyzed using different approaches: amplicon sequence variants and operational taxonomic units from the 16S rRNA gene amplicons and operational taxonomic units from the 16S rRNA genes of the metagenomes. We found that prokaryotic rare biosphere communities are specific to certain water masses, and that the majority of the rare taxa identified were always rare and disappeared in at least one sample under changing conditions, suggesting their high sensitivity to environmental heterogeneity. In addition, our methodological comparison revealed a good performance of 16S rRNA gene amplicon sequencing in describing rare biosphere patterns, while the metagenome-derived data were better to capture a significant diversity of so-far uncultivated rare taxa. Our analysis on the dynamics of the rare prokaryotic biosphere, by combining different methodological approaches, improves the description of the types of rarity predicted from Community Assembly theory in the Arctic Ocean. UIDB/04423/2020; UIDP/04565/2020; Programa Operacional Regional de Lisboa 007317 info:eu-repo/semantics/publishedVersion

Polyketide synthases (PKSs) and non-ribosomal peptide synthetases (NRPSs) are mega enzymes responsible for the biosynthesis of a large fraction of natural products (NPs). Molecular markers for biosynthetic genes, such as the ketosynthase (KS) domain of PKSs, have been used to assess the diversity and distribution of biosynthetic genes in complex microbial communities. More recently, metagenomic studies have complemented and enhanced this approach by allowing the recovery of complete biosynthetic gene clusters (BGCs) from environmental DNA. In this study, the distribution and diversity of biosynthetic genes and clusters from Arctic Ocean samples (NICE-2015 expedition), was assessed using PCR-based strategies coupled with high-throughput sequencing and metagenomic analysis. In total, 149 KS domain OTU sequences were recovered, 36 % of which could not be assigned to any known BGC. In addition, 74 bacterial metagenome-assembled genomes were recovered, from which 179 BGCs were extracted. A network analysis identified potential new NP families, including non-ribosomal peptides and polyketides. Complete or near-complete BGCs were recovered, which will enable future heterologous expression efforts to uncover the respective NPs. Our study represents the first report of biosynthetic diversity assessed for Arctic Ocean metagenomes and highlights the potential of Arctic Ocean planktonic microbiomes for the discovery of novel secondary metabolites. The strategy employed in this study will enable future bioprospection, by identifying promising samples for bacterial isolation efforts, while providing also full-length BGCs for heterologous expression.

We investigated diets of 24 Barents Sea zooplankton taxa to understand pelagic food-web processes during late summer, including the importance of sea ice algae-produced carbon. This was achieved by combining insights derived from multiple and complementary trophic marker approaches to construct individual aspects of feeding. Specifically, we determined proportions of algal-produced fatty acids (FAs) to reflect the reliance on diatom- versus dinoflagellate-derived carbon, highly branched isoprenoid (HBI) lipids that distinguish between ice-associated and pelagic carbon sources, and sterols to indicate the degree of carnivory. Copepods had the strongest diatom signal based on FAs, while a lack of sea ice algae-associated HBIs (IP25, IPSO25) suggested that they fed on pelagic rather than ice-associated diatoms. The amphipodThemisto libellulaand the ctenophoresBeroë cucumisandMertensia ovumhad a higher contribution of dinoflagellate-produced FAs. There was a high degree of carnivory in this food web, as indicated by the FA carnivory index 18:1(n−9)/18:1(n−7) (mean value < 1 only in the pteropodClione limacina), the presence of copepod-associated FAs in most of the taxa, and the absence of algal-produced HBIs in small copepod taxa, such asOithona similisandPseudocalanusspp. The coherence between concentrations of HBIs and phytosterols within individuals suggested that phytosterols provide a good additional indication for algal ingestion. Sea ice algae-associated HBIs were detected in six zooplankton species (occurring in krill, amphipods, pteropods, and appendicularians), indicating an overall low to moderate contribution of ice-associated carbon from late-summer sea ice to pelagic consumption. The unexpected occurrence of ice-derived HBIs in pteropods and appendicularians, however, suggests an importance of sedimenting ice-derived material at least for filter feeders within the water column at this time of year.

The growth of phytoplankton at high latitudes was generally thought to begin in open waters of the marginal ice zone once the highly reflective sea ice retreats in spring, solar elevation increases, and surface waters become stratified by the addition of sea-ice melt water. In fact, virtually all recent large-scale estimates of primary production in the Arctic Ocean (AO) assume that phytoplankton production in the water column under sea ice is negligible. However, over the past two decades, an emerging literature showing significant under-ice phytoplankton production on a pan-Arctic scale has challenged our paradigms of Arctic phytoplankton ecology and phenology. This evidence, which builds on previous, but scarce reports, requires the Arctic scientific community to change its perception of traditional AO phenology and urgently revise it. In particular, it is essential to better comprehend, on small and large scales, the changing and variable icescapes, the under-ice light field and biogeochemical cycles during the transition from sea-ice covered to ice-free Arctic waters. Here, we provide a baseline of our current knowledge of under-ice blooms (UIBs), by defining their ecology and their environmental setting, but also their regional peculiarities (in terms of occurrence, magnitude, and assemblages), which is shaped by a complex AO. To this end, a multidisciplinary approach, i.e., combining expeditions and modern autonomous technologies, satellite, and modeling analyses, has been used to provide an overview of this pan-Arctic phenological feature, which will become increasingly important in future marine Arctic biogeochemical cycles.

The attenuation coefficient b is one of the most common ways to describe how strong the carbon flux is attenuated throughout the water column. Therefore, b is an essential input variable in many carbon flux and climate models. Marsay et al. (2015, https://doi.org/10.1073/pnas.1415311112) proposed that the median surface water temperature (0–500 m) may be a predictor of b, but our observations from Arctic waters challenge this hypothesis. We found a highly variable attenuation coefficient (b = 0.43–1.84) in cold Arctic waters (

The genesis of phytoplankton blooms and the fate of their biomass in iron-limited, high-nutrient−low-chlorophyll regions can be studied under natural conditions with ocean iron fertilization (OIF) experiments. The Indo-German OIF experiment LOHAFEX was carried out over 40 d in late summer 2009 within the cold core of a mesoscale eddy in the productive southwest Atlantic sector of the Southern Ocean. Silicate concentrations were very low, and phytoplankton biomass was dominated by autotrophic nanoflagellates (ANF) in the size range 3−10 μm. As in all previous OIF experiments, the phytoplankton responded to iron fertilization by increasing the maximum quantum yield (Fv/Fm) and cellular chlorophyll levels. Within 3 wk, chlorophyll levels tripled and ANF biomass doubled. With the exception of some diatoms and dinoflagellates, the biomass levels of all other groups of the phyto- and protozooplankton (heterotrophic nanoflagellates, dinoflagellates and ciliates) remained remarkably stable throughout the experiment both inside and outside the fertilized patch. We attribute the unusually high biomass attained and maintained by ANF to the absence of their grazers, the salps, and to constraints on protozooplankton grazers by heavy predation exerted by the large copepod stock. The resistance to change of the ecosystem structure over 38 d after fertilization, indicated by homogeneity at regional and temporal scales, suggests that it was locked into a stable, mature state that had evolved in the course of the seasonal cycle. The LOHAFEX bloom provides a case study of a resistant/robust dynamic equilibrium between auto- and heterotrophic ecosystem components resulting in low vertical flux both inside and outside the patch despite high biomass levels.

The LOHAFEX iron fertilization experiment was conducted for 39 days in the closed core of a cyclonic mesoscale eddy located along the Antarctic Polar Front in the Atlantic sector of the Southern Ocean. Mixed layer (ML) waters were characterized by high nitrate (~ 20 μM), low dissolved iron (DFe ~ 0.2 nM) and low silicate concentrations (below 1 μM) restricting diatom growth. Upon initial fertilization, chlorophyll-a doubled during the first two weeks and stabilized thereafter, despite a second fertilization on day 21, due to an increase in grazing pressure. Biomass at the different trophic levels was mostly comprised of small autotrophic flagellates, the large copepod Calanus simillimus and the amphipod Themisto gaudichaudii. The downward flux of particulate material comprised mainly copepod fecal pellets that were remineralized in the upper 150 m of the water column with no significant deeper export. DFe concentrations in the upper 200 m were not significantly affected by the two fertilizations but after day 14 showed a greater variability (ranging from 0.3 to 1.3 nM) without a clear vertical pattern. Particulate iron concentrations (measured after 2 months at pH 1.4) decreased with time and showed a vertical pattern that indicated an important non-biogenic component at the bottom of the mixed layer. In order to assess the contribution of copepod grazing to iron cycling we used two different approaches: first, we measured for the first time in a field experiment copepod fecal pellet concentrations in the water column together with the iron content per pellet, and second, we devised a novel analytical scheme based on a two-step leaching protocol to estimate the contribution of copepod fecal pellets to particulate iron in the water column. Analysis of the iron content of isolated fecal pellets from C. simillimus showed that after the second fertilization, the iron content per fecal pellet was ~ 5 fold higher if the copepod had been captured in fertilized waters. We defined a new fraction termed leachable iron (pH 2.0) in 48 h (LFe48h) that, for the conditions during LOHAFEX, was shown to be an excellent proxy for the concentration of iron contained in copepod fecal pellets. We observed that, as a result of the second fertilization, iron accumulated in copepod fecal pellets and remained high at one third of the total iron stock in the upper 80 m. We hypothesize that our observations are due to a combination of two biological processes. First, phagotrophy of iron colloids freshly formed after the second fertilization by the predominant flagellate community resulted in higher Fe:C ratios per cell that, via grazing, lead to iron enrichment in copepod fecal pellets in fertilized waters. Second, copepod coprophagy could explain the rapid recycling of particulate iron in the upper 100–150 m, the accumulation of LFe48h in the upper 80 m after the second fertilization and provided the iron required for the maintenance of the LOHAFEX bloom for many weeks. Our results provide the first quantitative evidence of the major ecological relevance of copepods and their fecal products in the cycling of iron in silicate depleted areas of the Southern Ocean., This work was supported by the Government of Spain (MINECO CTM2008-01864-E/ANT and CTM2014-59244-C3-3-R). LML was supported by the Campus de Excelencia Internacional Program (Ministerio de Educación, Cultura y Deportes, España).

We report the identification of a range of C 25 highly branched isoprenoid (HBI) alkenes and certain sterols in filtered phytoplankton samples obtained from western Svalbard (Arctic) and near South Georgia (South Atlantic, sub-Antarctic) in 2016 and 2014, respectively. The C 25 HBIs contained 3–5 double bonds and had structures identified previously from analysis of laboratory diatom cultures. The same HBIs were also identified in individual diatom taxa isolated from the mixed assemblages and with reasonably similar distributions. Thus, C 25 HBIs were identified in Rhizosolenia setigera isolated from western Svalbard near-surface waters, while the same HBIs were also found in R. polydactyla f. polydactyla and R. hebetata f. semispina picked from seawater collected from a site in the South Atlantic. The main sterol composition was slightly different between the two locations, with cholesta-5,24-dien-3β-ol (desmosterol) identified as one of the major components in the sample from West Svalbard, consistent with the diatom assemblage being dominated by R. setigera . In contrast, the major sterol in the South Atlantic sample was cholesta-5,22-dien-3β-ol (22-dehydrocholesterol), likely reflecting the relatively high proportion of the genus Pseudo-nitzschia . For both locations, the suite of HBIs included a tri-unsaturated isomer (HBI III; 6 Z -2,6,10,14-tetramethyl-9-(3′-methylpent-4-enylidene)-pentadec-6-ene), proposed in previous studies as a potential proxy measure of pelagic sea ice-edge conditions, and thus a counterpart to the mono- and di-unsaturated HBIs IP 25 and IPSO 25 , which have been used as seasonal sea ice proxies in the Arctic and Antarctic, respectively. HBI III has been reported previously in sediments from West Svalbard and we report here its occurrence in a small number of surface sediments from the South Atlantic. For both regions, HBI III was present as one of the major HBIs in sediments, which contrasts with the HBI distributions in the filtered phytoplankton samples, where HBIs with four and five double bonds were the major components. Differences in HBI distributions between phytoplankton and sediment samples may potentially be due to the presence of other (unanalysed) diatoms in the filtered water samples, seasonal/annual variability in the production of HBIs by a range of diatoms, differential degradation of HBIs between sources and sediments, or a combination of these. Interestingly, we did not detect any C 30 HBIs in the water samples, picked cells or sediments from either location, despite earlier reports of these lipids in laboratory cultures of R. setigera . This study represents the first source identification of certain C 25 HBI lipids under in situ pelagic conditions.

Large changes in the sea ice regime of the Arctic Ocean have occurred over the last decades justifying the development of models to forecast sea ice physics and biogeochemistry. The main goal of this study is to evaluate the performance of the Los Alamos Sea Ice Model (CICE) to simulate physical and biogeochemical properties at time scales of a few weeks and to use the model to analyze ice algal bloom dynamics in different types of ice. Ocean and atmospheric forcing data and observations of the evolution of the sea ice properties collected from 18 April to 4 June 2015, during the Norwegian young sea ICE expedition, were used to test the CICE model. Our results show the following: (i) model performance is reasonable for sea ice thickness and bulk salinity; good for vertically resolved temperature, vertically averaged Chl a concentrations, and standing stocks; and poor for vertically resolved Chl a concentrations. (ii) Improving current knowledge about nutrient exchanges, ice algal recruitment, and motion is critical to improve sea ice biogeochemical modeling. (iii) Ice algae may bloom despite some degree of basal melting. (iv) Ice algal motility driven by gradients in limiting factors is a plausible mechanism to explain their vertical distribution. (v) Different ice algal bloom and net primary production (NPP) patterns were identified in the ice types studied, suggesting that ice algal maximal growth rates will increase, while sea ice vertically integrated NPP and biomass will decrease as a result of the predictable increase in the area covered by refrozen leads in the Arctic Ocean.

During the Norwegian young sea ICE expedition (N-ICE2015) from January to June 2015 the pack ice in the Arctic Ocean north of Svalbard was studied during four drifts between 83° and 80° N. This pack ice consisted of a mix of second-year, first-year and young ice. The physical properties and ice algal community composition was investigated in the three different ice types during the winter-spring-summer transition. Our results indicate that algae remaining in sea ice that survived the summer melt season are subsequently trapped in the upper layers of the ice column during winter and may function as an algal seed repository. Once the connectivity in the entire ice column is established, as a result of temperature-driven increase in ice porosity during spring, algae in the upper parts of the ice are able to migrate towards the bottom and initiate the ice-algal spring bloom. Furthermore, this algal repository might seed the bloom in younger ice formed in adjacent leads. This mechanism was studied in detail for the often dominating ice diatom Nitzschia frigida.The proposed seeding mechanism may be compromised due to the disappearance of older ice in the anticipated regime shift towards a seasonally ice-free Arctic Ocean.

We performed measurements of carbon dioxide fugacity (fCO2) in the surface water under Arctic sea ice from January to June 2015 during the Norwegian young sea ICE (N-ICE2015) expedition. Over this period, the ship drifted with four different ice floes and covered the deep Nansen Basin, the slopes north of Svalbard and the Yermak Plateau. This unique winter-to-spring dataset includes the first winter-time under-ice water fCO2 observations in this region. The observed under-ice fCO2 ranged between 315 µatm in winter and 153 µatm in spring, hence was undersaturated relative to the atmospheric fCO2. Although the sea ice partly prevented direct CO2 exchange between ocean and atmosphere, frequently occurring leads and breakup of the ice sheet promoted sea-air CO2 fluxes. The CO2 sink varied between 0.3 and 86 mmol C m−2 d−1, depending strongly on the open-water fractions (OW) and storm events. The maximum sea-air CO2 fluxes occurred during storm events in February and June. In winter, the main drivers of the change in under-ice water fCO2 were dissolution of CaCO3 (ikaite) and vertical mixing. In June, in addition to these processes, primary production and sea-air CO2 fluxes were important. The cumulative loss due to CaCO3 dissolution of 0.7 mol C m−2 in the upper 10 m played a major role in sustaining the undersaturation of fCO2 during the entire study. The relative effects of the total fCO2 change due to CaCO3 dissolution was 38%, primary production 26%, vertical mixing 16%, sea-air CO2 fluxes 16%, and temperature and salinity insignificant. This article is protected by copyright. All rights reserved.

In spring 2015, we observed an extensive phytoplankton bloom of Phaeocystis pouchetii, with chlorophyll a concentrations up to 7.5 mg m−3, under compact snow-covered Arctic sea ice at 80-81˚N during the Norwegian young sea ICE (N-ICE2015) expedition. We investigated the influence of the under-ice bloom on inherent optical properties (IOPs) of the upper ocean. Absorption and scattering in the upper 20 m of the water column at visible wavebands increased threefold and tenfold, respectively, relative to pre-bloom conditions. The scattering-to-absorption ratio during the Phaeocystis under-ice bloom was higher than in previous Arctic studies investigating diatom blooms. During the bloom, absorption by colored dissolved organic matter (at 375 nm), seemingly of autochthonous origin, doubled. Total absorption by particles (at 440 nm), dominated by phytoplankton (> 90%), increased tenfold. Measured absorption and scattering in the water were used as inputs for a 1D coupled atmosphere-ice-ocean radiative transfer model (AccuRT) to investigate effects of altered IOPs on the under-ice light field. Multiple scattering between sea ice and phytoplankton in the ocean led to an increase in scalar irradiance in the photosynthetically active radiation range (Eo(PAR)) at the ice-ocean interface by 6–7% compared to pre-bloom situation. This increase could have a positive feedback on ice-algal and under-ice phytoplankton productivity. The ratio between Eo(PAR) and downwelling planar irradiance (Ed(PAR)) below sea ice reached 1.85. Therefore, the use of Ed(PAR) might significantly underestimate the amount of PAR available for photosynthesis underneath sea ice. Our findings could help to improve light parameterizations in primary production models.

The Arctic Ocean is rapidly changing from thicker multiyear to thinner first-year ice cover, with significant consequences for radiative transfer through the ice pack and light availability for algal growth. A thinner, more dynamic ice cover will possibly result in more frequent leads, covered by newly formed ice with little snow cover. We studied a refrozen lead (≤0.27 m ice) in drifting pack ice north of Svalbard (80.5–82.4 °N) in May-June 2015 during the Norwegian young sea ICE expedition (N-ICE2015). We measured downwelling incident and ice transmitted spectral irradiance, and colored dissolved organic matter (CDOM), particle absorption, ultraviolet (UV)-protecting mycosporine-like amino acids (MAAs) and chlorophyll a (Chl a) in melted sea ice samples. We found occasionally very high MAA concentrations (up to 39 mg m-3, mean 4.5 ± 7.8 mg m-3) and MAA to Chl a ratios (up to 6.3, mean 1.2 ± 1.3). Disagreement in modelled and observed transmittance in the UV range let us conclude that MAA signatures in CDOM absorption spectra may be artefacts due to osmotic shock during ice melting. Although observed PAR transmittance through the thin ice was 5–40 times that of the adjacent thicker ice with deep snow cover, ice algal standing stocks were low (≤2.31 mg Chl a m-2) and similar to the adjacent ice. Ice algal accumulation in the lead was possibly delayed by the low inoculum and the time needed for photoacclimation to the high-light environment. However, leads are important for phytoplankton growth by acting like windows into the water column.

A number of common sterols were identified in sea ice diatoms from the Arctic and the Antarctic. The main sterols in floating sea ice algal aggregates collected from Resolute Passage (Canadian Arctic) and Nansen Basin (North Svalbard) in 2012 were 22E-dehydrocholesterol, cholesterol, epi-brassicasterol, 24-methylenecholesterol and 24-ethylcholesterol, although the distribution varied between the two locations, likely reflecting compositional differences in diatom taxa. The three major sterols in cells of Berkeleya adeliensis picked from a melted sea ice core collected from Ryder Bay in the Antarctic Peninsula in 2014, were 24-ethylcholesterol, cholesterol and 22E-dehydrocholesterol. We suggest that certain sea ice diatoms may thus contribute to the sedimentary budget of common sterols in seasonally sea ice-covered locations following ice melt.

Recent observations in the Atlantic sector of the Arctic reveal a sea-ice scape that functions quite differently than earlier assumed. It is also in a state where it responds more quickly to external forcing. Significant thinning of sea ice in this region has resulted in emergence of new features of the ice pack, that in many instances resemble the seasonal ice pack in the Southern Ocean. Thus in some sense an “antarctification” of the sea ice in the Arctic has occurred. This is a consequence of a thinner sea ice cover and a heavy enough snow load that now results in features, such as flooding- the phenomenon not thought to widely occur in the Arctic before. In addition to this, the thinner ice pack is more dynamic resulting in more deformation of the ice pack. These emerging physical properties of the new icescape also impact the functioning of the ice-associated ecosystem.

Phytoplankton phenology is a key driver of biological and chemical processes in marine ecosystems because it directly affects cycling of nutrients, the strength of the biological carbon pump, and energy transfer to higher tropic levels. However, phytoplankton time-series from the Arctic are scant, thus limiting our ability to link phytoplankton phenology to environmental variability. Kongsfjorden on the west coast of Spitsbergen is an established coastal monitoring site at the entrance to the Arctic Ocean. In this review we have compiled previously published phytoplankton investigations, chlorophyll fluorescence time-series data and unpublished phytoplankton data covering the years 2002–2014 from Kongsfjorden and the shelf outside the fjord to elaborate the most pertinent environmental factors responsible for the seasonal and inter-annual variability in phytoplankton bloom dynamics, biomass and species composition. In general, phytoplankton dynamics in Kongsfjorden follow the classic spring-bloom paradigm, with the main biomass peak in April–May dominated by spore-forming diatom species and the colony-forming haptophyte Phaeocystis pouchetii, followed by a diverse, but low biomass community characterised by dinoflagellates and small flagellates and their protozoan grazers during summer. Despite this general trend, phytoplankton phenology is subject to large inter-annual variability with no clear long-term trend. This variability can be mainly attributed to variability in the magnitude and depth of Atlantic Water (AW) inflow, sea ice cover and glacier melt-water discharge. We have shown the impact of environmental variability on phytoplankton phenology, but high-resolution monitoring of annual cycles over many years is required to resolve the ephemeral variations of phytoplankton populations in space and time against the backdrop of climate change.

Published version, licensed by the CC BY 4.0, available at http://dx.doi.org/10.1038/s41598-019-45574-5 A large retreat of sea-ice in the ‘stormy’ Atlantic Sector of the Arctic Ocean has become evident through a series of record minima for the winter maximum sea-ice extent since 2015. Results from the Norwegian young sea ICE (N-ICE2015) expedition, a five-month-long (Jan-Jun) drifting ice station in first and second year pack-ice north of Svalbard, showcase how sea-ice in this region is frequently affected by passing winter storms. Here we synthesise the interdisciplinary N-ICE2015 dataset, including independent observations of the atmosphere, snow, sea-ice, ocean, and ecosystem. We build upon recent results and illustrate the different mechanisms through which winter storms impact the coupled Arctic sea-ice system. These short-lived and episodic synoptic-scale events transport pulses of heat and moisture into the Arctic, which temporarily reduce radiative cooling and henceforth ice growth. Cumulative snowfall from each sequential storm deepens the snow pack and insulates the sea-ice, further inhibiting ice growth throughout the remaining winter season. Strong winds fracture the ice cover, enhance ocean-ice-atmosphere heat fluxes, and make the ice more susceptible to lateral melt. In conclusion, the legacy of Arctic winter storms for sea-ice and the ice-associated ecosystem in the Atlantic Sector lasts far beyond their short lifespan.

Marine ecosystems at high latitudes are characterized by extreme seasonal changes in light conditions, as well as a limited period of high primary production during spring and early summer. As light returns at the end of winter to Arctic ice-covered seas, a first algal bloom takes place in the bottom layer of the sea ice. This bottom ice algae community develops through three distinct phases in the transition from winter to spring, starting with phase I, a predominantly net heterotroph community that has limited interaction with the pelagic or benthic realms. Phase II begins in the spring once light for photosynthesis becomes available at the ice bottom, although interaction with the water column and benthos remains limited. The transition to the final phase III is then mainly driven by a balance of atmospheric and oceanographic forcing that induce structural changes in the sea ice and ultimately the removal of algal biomass from the ice. Due to limited data availability an incomplete understanding exists of all the processes determining ice algal bloom phenology and the considerable geographic differences in sympagic algal standing stocks and primary production. We present here the first pan-Arctic compilation of available time-series data on vernal sea ice algal bloom development and identify the most important factors controlling its development and termination. Using data from the area surrounding Resolute Bay (Nunavut, Canada) as an example, we support previous investigations that snow cover on top of the ice influences sea ice algal phenology, with highest biomass development, but also earliest termination of blooms, under low snow cover. We also provide a pan-Arctic overview of sea ice algae standing stocks and primary production, and discuss the pertinent processes behind the geographic differences we observed. Finally, we assess potential future changes in vernal algal bloom phenology as a consequence of climate change, including their importance to different groups of grazers.

The Marginal Ice Zone (MIZ) of the Arctic Ocean is changing rapidly due to a warming Arctic climate with commensurate reductions in sea ice extent and thickness. This Pan-Arctic review summarizes the main changes in the Arctic ocean–sea ice–atmosphere (OSA) interface, with implications for primary- and secondary producers in the ice and the underlying water column. Changes in the Arctic MIZ were interpreted for the period 1979–2010, based on best-fit regressions for each month. Trends of increasingly open water were statistically significant for each month, with quadratic fit for August–November, illustrating particularly strong seasonal feedbacks in sea-ice formation and decay. Geographic interpretations of physical and biological changes were based on comparison of regions with significant changes in sea ice: (1) The Pacific Sector of the Arctic Ocean including the Canada Basin and the Beaufort, Chukchi and East Siberian seas; (2) The Canadian Arctic Archipelago; (3) Baffin Bay and Hudson Bay; and (4) the Barents and Kara seas. Changes in ice conditions in the Barents sea/Kara sea region appear to be primarily forced by ocean heat fluxes during winter, whereas changes in the other sectors appear to be more summer–autumn related and primarily atmospherically forced. Effects of seasonal and regional changes in OSA-system with regard to increased open water were summarized for photosynthetically available radiation, nutrient delivery to the euphotic zone, primary production of ice algae and phytoplankton, ice-associated fauna and zooplankton, and gas exchange of CO2. Changes in the physical factors varied amongst regions, and showed direct effects on organisms linked to sea ice. Zooplankton species appear to be more flexible and likely able to adapt to variability in the onset of primary production. The major changes identified for the ice-associated ecosystem are with regard to production timing and abundance or biomass of ice flora and fauna, which are related to regional changes in sea-ice conditions.

One of the most prominent manifestations of climate change is the changing Arctic sea-ice regime with a reduction in the summer sea-ice extent and a shift from thicker, perennial multiyear ice towards thinner, first-year ice. These changes in the physical environment are likely to impact microbial communities, a key component of Arctic marine food webs and biogeochemical cycles. During the Norwegian young sea ICE expedition (N-ICE2015) north of Svalbard, seawater samples were collected at the surface (5 m), subsurface (20 or 50 m), and mesopelagic (250 m) depths on 9 March, 27 April, and 16 June 2015. In addition, several physical and biogeochemical data were recorded to contextualize the collected microbial communities. Through the massively parallel sequencing of the small subunit ribosomal RNA amplicon and metagenomic data, this work allows studying the Arctic’s microbial community structure during the late winter to early summer transition. Results showed that, at compositional level, Alpha- (30.7%) and Gammaproteobacteria (28.6%) are the most frequent taxa across the prokaryotic N-ICE2015 collection, and also the most phylogenetically diverse. Winter to early summer trends were quite evident since there was a high relative abundance of thaumarchaeotes in the under-ice water column in late winter while this group was nearly absent during early summer. Moreover, the emergence of Flavobacteria and the SAR92 clade in early summer might be associated with the degradation of a spring bloom of Phaeocystis. High relative abundance of hydrocarbonoclastic bacteria, particularly Alcanivorax (54.3%) and Marinobacter (6.3%), was also found. Richness showed different patterns along the depth gradient for prokaryotic (highest at mesopelagic depth) and protistan communities (higher at subsurface depths). The microbial N-ICE2015 collection analyzed in the present study provides comprehensive new knowledge about the pelagic microbiota below drifting Arctic sea-ice. The higher microbial diversity found in late winter/early spring communities reinforces the need to continue with further studies to properly characterize the winter microbial communities under the pack-ice.

The importance of newly formed sea ice in spring is likely to increase with formation of leads in a more dynamic Arctic icescape. We followed the ice algal species succession in young ice (≤ 0.27 m) in spring at high temporal resolution (sampling every second day for 1 month in May–June 2015) in the Arctic Ocean north of Svalbard. We document the early development of the ice algal community based on species abundance and chemotaxonomic marker pigments, and relate the young-ice algal community to the communities in the under-ice water column and the surrounding older ice. The seeding source seemed to vary between algal groups. Dinoflagellates were concluded to originate from the water column and diatoms from the surrounding older ice, which emphasizes the importance of older ice as a seeding source over deep oceanic regions and in early spring when algal abundance in the water column is low. In total, 120 taxa (80 identified to species or genus level) were recorded in the young ice. The protist community developed over the study period from a ciliate, flagellate, and dinoflagellate dominated community to one dominated by pennate diatoms. Environmental variables such as light were not a strong driver for the community composition, based on statistical analysis and comparison to the surrounding thicker ice with low light transmission. The photoprotective carotenoids to Chl a ratio increased over time to levels found in other high-light habitats, which shows that the algae were able to acclimate to the light levels of the thin ice. The development into a pennate diatom-dominated community, similar to the older ice, suggests that successional patterns tend toward ice-associated algae fairly independent of environmental conditions like light availability, season or ice type, and that biological traits, including morphological and physiological specialization to the sea ice habitat, play an important role in colonization of the sea ice environment. However, recruitment of ice-associated algae could be negatively affected by the ongoing loss of older ice, which acts as a seeding repository. Copyright © 2018 Kauko, Olsen, Duarte, Peeken, Granskog, Johnsen, Fernández-Méndez, Pavlov, Mundy and Assmy. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.