Philipp Assmy

Synthesis of iron fertilization experiments: From the Iron Age in the Age of Enlightenment

Comparison of eight iron experiments shows that maximum Chl a, the maximum DIC removal, and the overall DIC/Fe efficiency all scale inversely with depth of the wind mixed layer (WML) defining the light environment. Moreover, lateral patch dilution, sea surface irradiance, temperature, and grazing play additional roles. The Southern Ocean experiments were most influenced by very deep WMLs. In contrast, light conditions were most favorable during SEEDS and SERIES as well as during IronEx-2. The two extreme experiments, EisenEx and SEEDS, can be linked via EisenEx bottle incubations with shallower simulated WML depth. Large diatoms always benefit the most from Fe addition, where a remarkably small group of thriving diatom species is dominated by universal response of Pseudo-nitzschia spp. Significant response of these moderate (10–30 μm), medium (30–60 μm), and large (>60 μm) diatoms is consistent with growth physiology determined for single species in natural seawater. The minimum level of “dissolved” Fe (filtrate < 0.2 μm) maintained during an experiment determines the dominant diatom size class. However, this is further complicated by continuous transfer of original truly dissolved reduced Fe(II) into the colloidal pool, which may constitute some 75% of the “dissolved” pool. Depth integration of carbon inventory changes partly compensates the adverse effects of a deep WML due to its greater integration depths, decreasing the differences in responses between the eight experiments. About half of depth-integrated overall primary productivity is reflected in a decrease of DIC. The overall C/Fe efficiency of DIC uptake is DIC/Fe ∼ 5600 for all eight experiments. The increase of particulate organic carbon is about a quarter of the primary production, suggesting food web losses for the other three quarters. Replenishment of DIC by air/sea exchange tends to be a minor few percent of primary CO2 fixation but will continue well after observations have stopped. Export of carbon into deeper waters is difficult to assess and is until now firmly proven and quite modest in only two experiments.

The role of grazing in structuring Southern Ocean pelagic ecosystems and biogeochemical cycles

This review examines the links between pelagic ecology and ocean biogeochemistry with an emphasis on the role of the Southern Ocean in global cycling of carbon and silica. The structure and functioning of pelagic ecosystems is determined by the relationship between growth and mortality of its species populations. Whereas the key role of iron supply in conditioning the growth environment of land-remote oceans is now emerging, the factors shaping the mortality environment are still poorly understood. This paper addresses the role of grazing as a selective force operating on the structure and functioning of pelagic ecosystems within the larger conceptual framework of evolutionary ecology. That mortality due to grazing decreases with increasing cell size is widely taken for granted. We examine the impact of this principle across the range of size classes occupied by Southern Ocean plankton and show that relatively few species play crucial roles in the trophic structure and biogeochemical cycles of the Southern Ocean. Under iron-sufficient conditions, high growth rates of weakly silicified diatoms and Phaeocystis result in build-up of blooms that fuel “the food chain of the giants” (diatoms-krill-whales) and drive the carbon pump. In contrast, high grazing pressure of small copepods and salps on the regenerating microbial communities characteristic of the iron-limited Southern Ocean results in accumulation of large, heavily silicified diatoms that drive the silicon pump. The hypotheses we derive from field observations can be tested with in situ iron fertilization experiments.

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

Significance Silica-shelled diatoms dominate marine phytoplankton blooms and play a key role in ocean ecology and the global carbon cycle. We show how differences in ecological traits of dominant Southern Ocean diatom species, observed during the in situ European Iron Fertilization Experiment (EIFEX), can influence ocean carbon and silicon cycles. We argue that the ecology of thick-shelled diatom species, selected for by heavy copepod grazing, sequesters silicon relative to other nutrients in the deep Southern Ocean and underlying sediments to the detriment of diatom growth elsewhere. This evolutionary arms race provides a framework to link ecology with biogeochemistry of the ocean. 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.

Leads in Arctic pack ice enable early phytoplankton blooms below snow-covered sea ice

The Arctic icescape is rapidly transforming from a thicker multiyear ice cover to a thinner and largely seasonal first-year ice cover with significant consequences for Arctic primary production. One critical challenge is to understand how productivity will change within the next decades. Recent studies have reported extensive phytoplankton blooms beneath ponded sea ice during summer, indicating that satellite-based Arctic annual primary production estimates may be significantly underestimated. Here we present a unique time-series of a phytoplankton spring bloom observed beneath snow-covered Arctic pack ice. The bloom, dominated by the haptophyte algae Phaeocystis pouchetii, caused near depletion of the surface nitrate inventory and a decline in dissolved inorganic carbon by 16 ± 6 g C m−2. Ocean circulation characteristics in the area indicated that the bloom developed in situ despite the snow-covered sea ice. Leads in the dynamic ice cover provided added sunlight necessary to initiate and sustain the bloom. Phytoplankton blooms beneath snow-covered ice might become more common and widespread in the future Arctic Ocean with frequent lead formation due to thinner and more dynamic sea ice despite projected increases in high-Arctic snowfall. This could alter productivity, marine food webs and carbon sequestration in the Arctic Ocean.

Comparative molecular and morphological phylogenetic analyses of taxa in the Chaetocerotaceae (Bacillariophyta)

Kooistra W.H.F.C., Sarno D., Hernández-Becerril D.U., Assmy P., Di prisco C. and Montresor M. 2010. Comparative molecular and morphological phylogenetic analyses of taxa in the Chaetocerotaceae (Bacillariophyta). Phycologia 49: 471–500. DOI: 10.2216/09-59.1 The diatom family Chaetocerotaceae (Mediophyceae) includes two exclusively phytoplanktonic genera: Chaetoceros and Bacteriastrum. Its hallmark feature constitutes setae: hollow, spine-like appendages protruding from the valves. Chaetoceros is morphologically diverse, includes c. 400 described species and is common worldwide; whereas, Bacteriastrum includes only 11, is less common and occurs mainly in temperate and tropical seas. In the present study we gathered morphological information and/or sequence data from 86 strains belonging to 17 morphologically defined species in Chaetoceros and one in Bacteriastrum (B. cf. hyalinum). The Chaetoceros species included in this study belong to 14 of the 22 sections and two of the three subgenera: Chaetoceros (Phaeoceros) and Hyalochaete. A consensus cladogram reconstructed from states associated with morphological characters gathered from strains belonging to these 18 morphological taxa in the Chaetocerotaceae and Hemiaulus hauckii as outgroup resolved Bacteriastrum inside Chaetoceros and demonstrated monophyly for the subgenus Chaetoceros; whereas, the subgenus Hyalochaete was found to be paraphyletic. Molecular phylogenies inferred from the hypervariable region (D1–D4) of the LSU rRNA gene of all strains included in this study and from a subset of these strains corroborated the findings in the cladogram and showed evidence for cryptic or pseudocryptic diversity in C. curvisetus, C. debilis, C. diadema, C. lorenzianus, C. peruvianus and C. socialis. The molecular trees differed topologically from the morphological ones, but characters exhibiting only a few state changes in the molecular tree showed also only a few changes in the morphological one; whereas, characters showing multiple changes in the molecular tree revealed many changes in the morphological tree as well.

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.

Surface active substances in the upper water column during a Southern Ocean Iron Fertilization Experiment (EIFEX)

Surface active substances (SAS) in the water column were measured by voltammetry using the electrochemical probe o-nitrophenol (ONP) during EIFEX, a mesoscale open ocean iron enrichment experiment in the Southern Ocean. SAS levels were low throughout the experiment ( 0.02 mg L−1 Triton X-100 equivalents) were found at the end of the bloom particularly at density discontinuities where organic material may accumulate. Exudates from diatoms appeared to be the major source of SAS during EIFEX, either from direct extracellular release or in the action of being grazed upon by zooplankton.

Characterization of ikaite (CaCO3•6H2O) crystals in first year Arctic sea ice north of Svalbard

Abstract We identified ikaite crystals (CaCO3 ·6H2O) and examined their shape and size distribution in first-year Arctic pack ice, overlying snow and slush layers during the spring melt onset north of Svalbard. Additional measurements of total alkalinity (TA) were made for melted snow and sea-ice samples. Ikaite crystals were mainly found in the bottom of the snowpack, in slush and the surface layers of the sea ice where the temperature was generally lower and salinity higher than in the ice below. Image analysis showed that ikaite crystals were characterized by a roughly elliptical shape and a maximum caliper diameter of 201.0±115.9 μm (n = 918). Since the ice-melting season had already started, ikaite crystals may already have begun to dissolve, which might explain the lack of a relationship between ikaite crystal size and sea-ice parameters (temperature, salinity, and thickness of snow and ice). Comparisons of salinity and TA profiles for melted ice samples suggest that the precipitation/dissolution of ikaite crystals occurred at the top of the sea ice and the bottom of the snowpack during ice formation/melting processes.

Windows in Arctic sea ice: Light transmission and ice algae in a refrozen lead

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.