Benthic phosphorus cycling within the Eurasian marginal sea ice zone.

The Arctic Ocean region is currently undergoing dramatic changes, which will likely alter the nutrient cycles that underpin Arctic marine ecosystems. Phosphate is a key limiting nutrient for marine life but gaps in our understanding of the Arctic phosphorus (P) cycle persist. In this study, we investigate the benthic burial and recycling of phosphorus using sediments and pore waters from the Eurasian Arctic margin, including the Barents Sea slope and the Yermak Plateau. Our results highlight that P is generally lost from sediments with depth during organic matter respiration. On the Yermak Plateau, remobilization of P results in a diffusive flux of P to the seafloor of between 96 and 261 µmol m-2 yr-1. On the Barents Sea slope, diffusive fluxes of P are much larger (1736-2449 µmol m-2 yr-1), but these fluxes are into near-surface sediments rather than to the bottom waters. The difference in cycling on the Barents Sea slope is controlled by higher fluxes of fresh organic matter and active iron cycling. As changes in primary productivity, ocean circulation and glacial melt continue, benthic P cycling is likely to be altered with implications for P imported into the Arctic Ocean Basin. This article is part of the theme issue 'The changing Arctic Ocean: consequences for biological communities, biogeochemical processes and ecosystem functioning'.

Deep carbon export from a Southern Ocean iron-fertilized diatom bloom.

Fertilization of the ocean by adding iron compounds has induced diatom-dominated phytoplankton blooms accompanied by considerable carbon dioxide drawdown in the ocean surface layer. However, because the fate of bloom biomass could not be adequately resolved in these experiments, the timescales of carbon sequestration from the atmosphere are uncertain. Here we report the results of a five-week experiment carried out in the closed core of a vertically coherent, mesoscale eddy of the Antarctic Circumpolar Current, during which we tracked sinking particles from the surface to the deep-sea floor. A large diatom bloom peaked in the fourth week after fertilization. This was followed by mass mortality of several diatom species that formed rapidly sinking, mucilaginous aggregates of entangled cells and chains. Taken together, multiple lines of evidence-although each with important uncertainties-lead us to conclude that at least half the bloom biomass sank far below a depth of 1,000 metres and that a substantial portion is likely to have reached the sea floor. Thus, iron-fertilized diatom blooms may sequester carbon for timescales of centuries in ocean bottom water and for longer in the sediments.

Thick-shelled, grazer-protected diatoms decouple ocean carbon and silicon cycles in the iron-limited Antarctic Circumpolar Current.

Diatoms of the iron-replete continental margins and North Atlantic are key exporters of organic carbon. In contrast, diatoms of the iron-limited Antarctic Circumpolar Current sequester silicon, but comparatively little carbon, in the underlying deep ocean and sediments. Because the Southern Ocean is the major hub of oceanic nutrient distribution, selective silicon sequestration there limits diatom blooms elsewhere and consequently the biotic carbon sequestration potential of the entire ocean. We investigated this paradox in an in situ iron fertilization experiment by comparing accumulation and sinking of diatom populations inside and outside the iron-fertilized patch over 5 wk. A bloom comprising various thin- and thick-shelled diatom species developed inside the patch despite the presence of large grazer populations. After the third week, most of the thinner-shelled diatom species underwent mass mortality, formed large, mucous aggregates, and sank out en masse (carbon sinkers). In contrast, thicker-shelled species, in particular Fragilariopsis kerguelensis, persisted in the surface layers, sank mainly empty shells continuously, and reduced silicate concentrations to similar levels both inside and outside the patch (silica sinkers). These patterns imply that thick-shelled, hence grazer-protected, diatom species evolved in response to heavy copepod grazing pressure in the presence of an abundant silicate supply. The ecology of these silica-sinking species decouples silicon and carbon cycles in the iron-limited Southern Ocean, whereas carbon-sinking species, when stimulated by iron fertilization, export more carbon per silicon. Our results suggest that large-scale iron fertilization of the silicate-rich Southern Ocean will not change silicon sequestration but will add carbon to the sinking silica flux.

Release of ballast material during sea-ice melt enhances carbon export in the Arctic Ocean.

Globally, the most intense uptake of anthropogenic carbon dioxide (CO2) occurs in the Atlantic north of 50°N, and it has been predicted that atmospheric CO2 sequestration in the Arctic Ocean will increase as a result of ice-melt and increased primary production. However, little is known about the impact of pan-Arctic sea-ice decline on carbon export processes. We investigated the potential ballasting effect of sea-ice derived material on settling aggregates and carbon export in the Fram Strait by combining 13 years of vertical flux measurements with benthic eDNA analysis, laboratory experiments, and tracked sea-ice distributions. We show that melting sea-ice in the Fram Strait releases cryogenic gypsum and terrigenous material, which ballasts sinking organic aggregates. As a result, settling velocities of aggregates increased ≤10-fold, resulting in ≤30% higher carbon export in the vicinity of the melting ice-edge. Cryogenic gypsum is formed in first-year sea-ice, which is predicted to increase as the Arctic is warming. Simultaneously, less sea-ice forms over the Arctic shelves, which is where terrigenous material is incorporated into sea-ice. Supporting this, we found that terrigenous fluxes from melting sea-ice in the Fram Strait decreased by >80% during our time-series. Our study suggests that terrigenous flux will eventually cease when enhanced sea-ice melt disrupts trans-Arctic sea-ice transport and thus, limit terrigenous-ballasted carbon flux. However, the predicted increase in Arctic primary production and gypsum formation may enhance gypsum-ballasted carbon flux and compensate for lowered terrigenous fluxes. It is thus unclear if sea-ice loss will reduce carbon export in the Arctic Ocean.

Wind-driven upwelling of iron sustains dense blooms and food webs in the eastern Weddell Gyre.

The Southern Ocean is a major sink of anthropogenic CO2 and an important foraging area for top trophic level consumers. However, iron limitation sets an upper limit to primary productivity. Here we report on a considerably dense late summer phytoplankton bloom spanning 9000 km2 in the open ocean of the eastern Weddell Gyre. Over its 2.5 months duration, the bloom accumulated up to 20 g C m-2 of organic matter, which is unusually high for Southern Ocean open waters. We show that, over 1997-2019, this open ocean bloom was likely driven by anomalies in easterly winds that push sea ice southwards and favor the upwelling of Warm Deep Water enriched in hydrothermal iron and, possibly, other iron sources. This recurring open ocean bloom likely facilitates enhanced carbon export and sustains high standing stocks of Antarctic krill, supporting feeding hot spots for marine birds and baleen whales.

Interactions between the ice algae Fragillariopsis cylindrus and microplastics in sea ice.

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.

Arctic warming interrupts the Transpolar Drift and affects long-range transport of sea ice and ice-rafted matter.

Sea ice is an important transport vehicle for gaseous, dissolved and particulate matter in the Arctic Ocean. Due to the recently observed acceleration in sea ice drift, it has been assumed that more matter is advected by the Transpolar Drift from shallow shelf waters to the central Arctic Ocean and beyond. However, this study provides first evidence that intensified melt in the marginal zones of the Arctic Ocean interrupts the transarctic conveyor belt and has led to a reduction of the survival rates of sea ice exported from the shallow Siberian shelves (-15% per decade). As a consequence, less and less ice formed in shallow water areas (<30 m) has reached Fram Strait (-17% per decade), and more ice and ice-rafted material is released in the northern Laptev Sea and central Arctic Ocean. Decreasing survival rates of first-year ice are visible all along the Russian shelves, but significant only in the Kara Sea, East Siberian Sea and western Laptev Sea. Identified changes affect biogeochemical fluxes and ecological processes in the central Arctic: A reduced long-range transport of sea ice alters transport and redistribution of climate relevant gases, and increases accumulation of sediments and contaminates in the central Arctic Ocean, with consequences for primary production, and the biodiversity of the Arctic Ocean.

A red tide in the pack ice of the Arctic Ocean.

In the Arctic Ocean ice algae constitute a key ecosystem component and the ice algal spring bloom a critical event in the annual production cycle. The bulk of ice algal biomass is usually found in the bottom few cm of the sea ice and dominated by pennate diatoms attached to the ice matrix. Here we report a red tide of the phototrophic ciliate Mesodinium rubrum located at the ice-water interface of newly formed pack ice of the high Arctic in early spring. These planktonic ciliates are not able to attach to the ice. Based on observations and theory of fluid dynamics, we propose that convection caused by brine rejection in growing sea ice enabled M. rubrum to bloom at the ice-water interface despite the relative flow between water and ice. We argue that red tides of M. rubrum are more likely to occur under the thinning Arctic sea ice regime.

High colored dissolved organic matter (CDOM) absorption in surface waters of the central-eastern Arctic Ocean: Implications for biogeochemistry and ocean color algorithms.

As consequences of global warming sea-ice shrinking, permafrost thawing and changes in fresh water and terrestrial material export have already been reported in the Arctic environment. These processes impact light penetration and primary production. To reach a better understanding of the current status and to provide accurate forecasts Arctic biogeochemical and physical parameters need to be extensively monitored. In this sense, bio-optical properties are useful to be measured due to the applicability of optical instrumentation to autonomous platforms, including satellites. This study characterizes the non-water absorbers and their coupling to hydrographic conditions in the poorly sampled surface waters of the central and eastern Arctic Ocean. Over the entire sampled area colored dissolved organic matter (CDOM) dominates the light absorption in surface waters. The distribution of CDOM, phytoplankton and non-algal particles absorption reproduces the hydrographic variability in this region of the Arctic Ocean which suggests a subdivision into five major bio-optical provinces: Laptev Sea Shelf, Laptev Sea, Central Arctic/Transpolar Drift, Beaufort Gyre and Eurasian/Nansen Basin. Evaluating ocean color algorithms commonly applied in the Arctic Ocean shows that global and regionally tuned empirical algorithms provide poor chlorophyll-a (Chl-a) estimates. The semi-analytical algorithms Generalized Inherent Optical Property model (GIOP) and Garver-Siegel-Maritorena (GSM), on the other hand, provide robust estimates of Chl-a and absorption of colored matter. Applying GSM with modifications proposed for the western Arctic Ocean produced reliable information on the absorption by colored matter, and specifically by CDOM. These findings highlight that only semi-analytical ocean color algorithms are able to identify with low uncertainty the distribution of the different optical water constituents in these high CDOM absorbing waters. In addition, a clustering of the Arctic Ocean into bio-optical provinces will help to develop and then select province-specific ocean color algorithms.

Arctic sea ice is an important temporal sink and means of transport for microplastic.

Microplastics (MP) are recognized as a growing environmental hazard and have been identified as far as the remote Polar Regions, with particularly high concentrations of microplastics in sea ice. Little is known regarding the horizontal variability of MP within sea ice and how the underlying water body affects MP composition during sea ice growth. Here we show that sea ice MP has no uniform polymer composition and that, depending on the growth region and drift paths of the sea ice, unique MP patterns can be observed in different sea ice horizons. Thus even in remote regions such as the Arctic Ocean, certain MP indicate the presence of localized sources. Increasing exploitation of Arctic resources will likely lead to a higher MP load in the Arctic sea ice and will enhance the release of MP in the areas of strong seasonal sea ice melt and the outflow gateways.

Inter-annual variability of transparent exopolymer particles in the Arctic Ocean reveals high sensitivity to ecosystem changes.

Transparent exopolymer particles (TEP) are a class of marine gel particles and important links between surface ocean biology and atmospheric processes. Derived from marine microorganisms, these particles can facilitate the biological pumping of carbon dioxide to the deep sea, or act as cloud condensation and ice nucleation particles in the atmosphere. Yet, environmental controls on TEP abundance in the ocean are poorly known. Here, we investigated some of these controls during the first multiyear time-series on TEP abundance for the Fram Strait, the Atlantic gateway to the Central Arctic Ocean. Data collected at the Long-Term Ecological Research observatory HAUSGARTEN during 2009 to 2014 indicate a strong biological control with highest abundance co-occurring with the prymnesiophyte Phaeocystis pouchetii. Higher occurrence of P. pouchetii in the Arctic Ocean has previously been related to northward advection of warmer Atlantic waters, which is expected to increase in the future. Our study highlights the role of plankton key species in driving climate relevant processes; thus, changes in plankton distribution need to be accounted for when estimating the ocean's biogeochemical response to global change.

Composition, buoyancy regulation and fate of ice algal aggregates in the Central Arctic Ocean.

Sea-ice diatoms are known to accumulate in large aggregates in and under sea ice and in melt ponds. There is recent evidence from the Arctic that such aggregates can contribute substantially to particle export when sinking from the ice. The role and regulation of microbial aggregation in the highly seasonal, nutrient- and light-limited Arctic sea-ice ecosystem is not well understood. To elucidate the mechanisms controlling the formation and export of algal aggregates from sea ice, we investigated samples taken in late summer 2011 and 2012, during two cruises to the Eurasian Basin of the Central Arctic Ocean. Spherical aggregates densely packed with pennate diatoms, as well as filamentous aggregates formed by Melosira arctica showed sign of different stages of degradation and physiological stoichiometries, with carbon to chlorophyll a ratios ranging from 110 to 66700, and carbon to nitrogen molar ratios of 8-35 and 9-40, respectively. Sub-ice algal aggregate densities ranged between 1 and 17 aggregates m(-2), maintaining an estimated net primary production of 0.4-40 mg C m(-2) d(-1), and accounted for 3-80% of total phototrophic biomass and up to 94% of local net primary production. A potential factor controlling the buoyancy of the aggregates was light intensity, regulating photosynthetic oxygen production and the amount of gas bubbles trapped within the mucous matrix, even at low ambient nutrient concentrations. Our data-set was used to evaluate the distribution and importance of Arctic algal aggregates as carbon source for pelagic and benthic communities.

Protist distribution in the Western Fram Strait in summer 2010 based on 454-pyrosequencing of 18S rDNA.

In this study, we present the first comprehensive analyses of the diversity and distribution of marine protist (micro-, nano-, and picoeukaryotes) in the Western Fram Strait, using 454-pyrosequencing and high-pressure liquid chromatography (HPLC) at five stations in summer 2010. Three stations (T1; T5; T7) were influenced by Polar Water, characterized by cold water with lower salinity (<33) and different extents of ice concentrations. Atlantic Water influenced the other two stations (T6; T9). While T6 was located in the mixed water zone characterized by cold water with intermediate salinity (~33) and high ice concentrations, T9 was located in warm water with high salinity (~35) and no ice-coverage at all. General trends in community structure according to prevailing environmental settings, observed with both methods, coincided well. At two stations, T1 and T7, characterized by lower ice concentrations, diatoms (Fragilariopsis sp., Porosira sp., Thalassiosira spp.) dominated the protist community. The third station (T5) was ice-covered, but has been ice-free for ~4 weeks prior to sampling. At this station, dinoflagellates (Dinophyceae 1, Woloszynskia sp. and Gyrodinium sp.) were dominant, reflecting a post-bloom situation. At station T6 and T9, the protist communities consisted mainly of picoeukaryotes, e.g., Micromonas spp. Based on our results, 454-pyrosequencing has proven to be an adequate tool to provide comprehensive information on the composition of protist communities. Furthermore, this study suggests that a snap-shot of a few, but well-chosen samples can provide an overview of community structure patterns and succession in a dynamic marine environment.

Export of algal biomass from the melting Arctic sea ice.

In the Arctic, under-ice primary production is limited to summer months and is restricted not only by ice thickness and snow cover but also by the stratification of the water column, which constrains nutrient supply for algal growth. Research Vessel Polarstern visited the ice-covered eastern-central basins between 82° to 89°N and 30° to 130°E in summer 2012, when Arctic sea ice declined to a record minimum. During this cruise, we observed a widespread deposition of ice algal biomass of on average 9 grams of carbon per square meter to the deep-sea floor of the central Arctic basins. Data from this cruise will contribute to assessing the effect of current climate change on Arctic productivity, biodiversity, and ecological function.

Floating ice-algal aggregates below melting arctic sea ice.

During two consecutive cruises to the Eastern Central Arctic in late summer 2012, we observed floating algal aggregates in the melt-water layer below and between melting ice floes of first-year pack ice. The macroscopic (1-15 cm in diameter) aggregates had a mucous consistency and were dominated by typical ice-associated pennate diatoms embedded within the mucous matrix. Aggregates maintained buoyancy and accumulated just above a strong pycnocline that separated meltwater and seawater layers. We were able, for the first time, to obtain quantitative abundance and biomass estimates of these aggregates. Although their biomass and production on a square metre basis was small compared to ice-algal blooms, the floating ice-algal aggregates supported high levels of biological activity on the scale of the individual aggregate. In addition they constituted a food source for the ice-associated fauna as revealed by pigments indicative of zooplankton grazing, high abundance of naked ciliates, and ice amphipods associated with them. During the Arctic melt season, these floating aggregates likely play an important ecological role in an otherwise impoverished near-surface sea ice environment. Our findings provide important observations and measurements of a unique aggregate-based habitat during the 2012 record sea ice minimum year.

Screening volatile organic compounds (VOCs) emissions from five marine phytoplankton species by head space gas chromatography/mass spectrometry (HS-GC/MS).

Five marine cosmopolitan phytoplankton species namely; Calcidiscus leptoporus, Emiliania huxleyi, Phaeodactylum tricornutum, Chaetoceros neogracilis and Dunaliella tertiolecta were screened for emissions of selected VOCs using head space gas chromatography/mass spectrometry (HS-GC/MS) in single ion mode. The VOCs investigated included isoprene and various halogenated compounds. Among the different algae groups, the two diatoms Ch. neogracilis and P. tricornutum were the strongest emitters of methyl bromide (CH3Br), and Ch. neogracilis was the strongest emitter of isoprene. Furthermore, we present evidence that several chlorinated organic compounds, normally considered as anthropogenic, can be produced from marine phytoplankton (namely chloroform, dichloromethane, trichloroethylene, tetrachloroethylene, chlorobenzene and dichlorobenzene).