Scarlett Trimborn

Iron Limitation Modulates Ocean Acidification Effects on Southern Ocean Phytoplankton Communities

The potential interactive effects of iron (Fe) limitation and Ocean Acidification in the Southern Ocean (SO) are largely unknown. Here we present results of a long-term incubation experiment investigating the combined effects of CO2 and Fe availability on natural phytoplankton assemblages from the Weddell Sea, Antarctica. Active Chl a fluorescence measurements revealed that we successfully cultured phytoplankton under both Fe-depleted and Fe-enriched conditions. Fe treatments had significant effects on photosynthetic efficiency (Fv/Fm; 0.3 for Fe-depleted and 0.5 for Fe-enriched conditions), non-photochemical quenching (NPQ), and relative electron transport rates (rETR). pCO2 treatments significantly affected NPQ and rETR, but had no effect on Fv/Fm. Under Fe limitation, increased pCO2 had no influence on C fixation whereas under Fe enrichment, primary production increased with increasing pCO2 levels. These CO2-dependent changes in productivity under Fe-enriched conditions were accompanied by a pronounced taxonomic shift from weakly to heavily silicified diatoms (i.e. from Pseudo-nitzschia sp. to Fragilariopsis sp.). Under Fe-depleted conditions, this functional shift was absent and thinly silicified species dominated all pCO2 treatments (Pseudo-nitzschia sp. and Synedropsis sp. for low and high pCO2, respectively). Our results suggest that Ocean Acidification could increase primary productivity and the abundance of heavily silicified, fast sinking diatoms in Fe-enriched areas, both potentially leading to a stimulation of the biological pump. Over much of the SO, however, Fe limitation could restrict this possible CO2 fertilization effect.

Transcriptome analyses reveal differential gene expression patterns between the life-cycle stages of Emiliania Huxleyi (haptophyta) and reflect specialization to different ecological niches

Coccolithophores, especially the abundant, cosmopolitan species Emiliania huxleyi (Lohmann) W. W. Hay et H. P. Mohler, are one of the main driving forces of the oceanic carbonate pump and contribute significantly to global carbon cycling, due to their ability to calcify. A recent study indicates that termination of diploid blooms by viral infection induces life‐cycle transition, and speculation has arisen about the role of the haploid, noncalcifying stage in coccolithophore ecology. To explore gene expression patterns in both life‐cycle stages, haploid and diploid cells of E. huxleyi (RCC 1217 and RCC 1216) were acclimated to limiting and saturating photon flux densities. Transcriptome analyses were performed to assess differential genomic expression related to different ploidy levels and acclimation light intensities. Analyses indicated that life‐cycle stages exhibit different properties of regulating genome expression (e.g., pronounced gene activation and gene silencing in the diploid stage), proteome maintenance (e.g., increased turnover of proteins in the haploid stage), as well as metabolic processing (e.g., pronounced primary metabolism and motility in the haploid stage and calcification in the diploid stage). Furthermore, higher abundances of transcripts related to endocytotic and digestive machinery were observed in the diploid stage. A qualitative feeding experiment indicated that both life‐cycle stages are capable of particle uptake (0.5 μm diameter) in late‐stationary growth phase. Results showed that the two life‐cycle stages represent functionally distinct entities that are evolutionarily shaped to thrive in the environment they typically inhabit.

Ocean acidification decreases the light-use efficiency in an Antarctic diatom under dynamic but not constant light

Summary There is increasing evidence that different light intensities strongly modulate the effects of ocean acidification (OA) on marine phytoplankton. The aim of the present study was to investigate interactive effects of OA and dynamic light, mimicking natural mixing regimes. The Antarctic diatom Chaetoceros debilis was grown under two pCO 2 (390 and 1000 μatm) and light conditions (constant and dynamic), the latter yielding the same integrated irradiance over the day. To characterize interactive effects between treatments, growth, elemental composition, primary production and photophysiology were investigated. Dynamic light reduced growth and strongly altered the effects of OA on primary production, being unaffected by elevated pCO 2 under constant light, yet significantly reduced under dynamic light. Interactive effects between OA and light were also observed for Chl production and particulate organic carbon quotas. Response patterns can be explained by changes in the cellular energetic balance. While the energy transfer efficiency from photochemistry to biomass production (Φe,C) was not affected by OA under constant light, it was drastically reduced under dynamic light. Contrasting responses under different light conditions need to be considered when making predictions regarding a more stratified and acidified future ocean.

Inorganic carbon utilization by Ross Sea phytoplankton across natural and experimental CO2 gradients.

We present results from a field study of inorganic carbon (C) acquisition by Ross Sea phytoplankton during Phaeocystis‐dominated early season blooms. Isotope disequilibrium experiments revealed that HCO3− was the primary inorganic C source for photosynthesis in all phytoplankton assemblages. From these experiments, we also derived relative enhancement factors for HCO3−/CO2 interconversion as a measure of extracellular carbonic anhydrase activity (eCA). The enhancement factors ranged from 1.0 (no apparent eCA activity) to 6.4, with an overall mean of 2.9. Additional eCA measurements, made using membrane inlet mass spectrometry (MIMS), yielded activities ranging from 2.4 to 6.9 U · [μg chl a]−1 (mean 4.1). Measurements of short‐term C‐fixation parameters revealed saturation kinetics with respect to external inorganic carbon, with a mean half‐saturation constant for inorganic carbon uptake (K1/2) of ∼380 μM. Comparison of our early springtime results with published data from late‐season Ross Sea assemblages showed that neither HCO3− utilization nor eCA activity was significantly correlated to ambient CO2 levels or phytoplankton taxonomic composition. We did, however, observe a strong negative relationship between surface water pCO2 and short‐term 14C‐fixation rates for the early season survey. Direct incubation experiments showed no statistically significant effects of pCO2 (10 to 80 Pa) on relative HCO3− utilization or eCA activity. Our results provide insight into the seasonal regulation of C uptake by Ross Sea phytoplankton across a range of pCO2 and phytoplankton taxonomic composition.

Southern Ocean phytoplankton physiology in a changing climate

The Southern Ocean (SO) is a major sink for anthropogenic atmospheric carbon dioxide (CO2), potentially harbouring even greater potential for additional sequestration of CO2 through enhanced phytoplankton productivity. In the SO, primary productivity is primarily driven by bottom up processes (physical and chemical conditions) which are spatially and temporally heterogeneous. Due to a paucity of trace metals (such as iron) and high variability in light, much of the SO is characterised by an ecological paradox of high macronutrient concentrations yet uncharacteristically low chlorophyll concentrations. It is expected that with increased anthropogenic CO2 emissions and the coincident warming, the major physical and chemical process that govern the SO will alter, influencing the biological capacity and functioning of the ecosystem. This review focuses on the SO primary producers and the bottom up processes that underpin their health and productivity. It looks at the major physico-chemical drivers of change in the SO, and based on current physiological knowledge, explores how these changes will likely manifest in phytoplankton, specifically, what are the physiological changes and floristic shifts that are likely to ensue and how this may translate into changes in the carbon sink capacity, net primary productivity and functionality of the SO.

Controls of primary production in two phytoplankton blooms in the Antarctic Circumpolar Current

The Antarctic Circumpolar Current has a high potential for primary production and carbon sequestration through the biological pump. In the current study, two large-scale blooms observed in 2012 during a cruise with R.V. Polarstern were investigated with respect to phytoplankton standing stocks, primary productivity and nutrient budgets. While net primary productivity was similar in both blooms, chlorophyll a –specific photosynthesis was more efficient in the bloom closer to the island of South Georgia (39 °W, 50 °S) compared to the open ocean bloom further east (12 °W, 51 °S). We did not find evidence for light being the driver of bloom dynamics as chlorophyll standing stocks up to 165 mg m−2 developed despite mixed layers as deep as 90 m. Since the two bloom regions differ in their distance to shelf areas, potential sources of iron vary. Nutrient (nitrate, phosphate, silicate) deficits were similar in both areas despite different bloom ages, but their ratios indicated more pronounced iron limitation at 12 °W compared to 39 °W. While primarily the supply of iron and not the availability of light seemed to control onset and duration of the blooms, higher grazing pressure could have exerted a stronger control toward the declining phase of the blooms.

The impact of iron limitation on the physiology of the Antarctic diatom Chaetoceros simplex

Iron availability strongly governs the growth of Southern Ocean phytoplankton. To investigate how iron limitation affects photosynthesis as well as the uptake of carbon and iron in the Antarctic diatom Chaetocerossimplex, a combination of chlorophyll a fluorescence measurements and radiotracer incubations in the presence and absence of chemical inhibitors was conducted. Iron limitation in C. simplex led to a decline in growth rates, photochemical efficiency and structural changes in photosystem II (PSII), including a reorganisation of photosynthetic units in PSII and an increase in size of the functional absorption cross section of PSII. Iron-limited cells further exhibited a reduced plastoquinone pool and decreased photosynthetic electron transport rate, while non-photochemical quenching and relative xanthophyll pigment content were strongly increased, suggesting a photoprotective response. Additionally, iron limitation resulted in a strong decline in carbon fixation and thus the particulate organic carbon quotas. Inhibitor studies demonstrated that, independent of the iron supply, carbon fixation was dependent on internal, but not on extracellular carbonic anhydrase activity. Orthovanadate more strongly inhibited iron uptake in iron-limited cells, indicating that P-type ATPase transporters are involved in iron uptake. The stronger reduction in iron uptake by ascorbate in iron-limited cells suggests that the re-oxidation of iron is required before it can be taken up and further supports the presence of a high-affinity iron transport pathway. The measured changes to photosystem architecture and shifts in carbon and iron uptake strategies in C. simplex as a result of iron limitation provide evidence for a complex interaction of these processes to balance the iron requirements for photosynthesis and carbon demand for sustained growth in iron-limited waters.

Two Southern Ocean diatoms are more sensitive to ocean acidification and changes in irradiance than the prymnesiophyte Phaeocystis antarctica

To better understand the impact of ocean acidification (OA) and changes in light availability on Southern Ocean phytoplankton physiology, we investigated the effects of pCO2 (380 and 800 µatm) in combination with low and high irradiance (20 or 50 and 200 µmol photons m-2  s-1 ) on growth, particulate organic carbon (POC) fixation and photophysiology in the three ecologically relevant species Chaetoceros debilis, Fragilariopsis kerguelensis and Phaeocystis antarctica. Irrespective of the light scenario, neither growth nor POC per cell was stimulated by OA in any of the tested species and the two diatoms even displayed negative responses in growth (e.g. C. debilis) or POC content (e.g. F. kerguelensis) under OA in conjunction with high light. For both diatoms, also maximum quantum yields of photosystem II (Fv /Fm ) were decreased under these conditions, indicating lowered photochemical efficiencies. To counteract the negative effects by OA and high light, the two diatoms showed diverging photoacclimation strategies. While cellular chlorophyll a (Chl a) and fucoxanthin contents were enhanced in C. debilis to potentially maximize light absorption, F. kerguelensis exhibited reduced Chl a per cell, increased disconnection of antennae from photosystem II reaction centers and strongly lowered absolute electron transport rates (ETR). The decline in ETRs in F. kerguelensis might be explained in terms of different species-specific strategies for tuning the available flux of adenosine triphosphate and nicotinamide adenine dinucleotide phosphate. Overall, our results revealed that P. antarctica was more tolerant to OA and changes in irradiance than the two diatoms, which may have important implications for biogeochemical cycling.

Meteorology and oceanography of the Atlantic sector of the Southern Ocean—a review of German achievements from the last decade

In the early 1980s, Germany started a new era of modern Antarctic research. The Alfred Wegener Institute Helmholtz Centre for Polar and Marine Research (AWI) was founded and important research platforms such as the German permanent station in Antarctica, today called Neumayer III, and the research icebreaker Polarstern were installed. The research primarily focused on the Atlantic sector of the Southern Ocean. In parallel, the German Research Foundation (Deutsche Forschungsgemeinschaft, DFG) started a priority program ‘Antarctic Research’ (since 2003 called SPP-1158) to foster and intensify the cooperation between scientists from different German universities and the AWI as well as other institutes involved in polar research. Here, we review the main findings in meteorology and oceanography of the last decade, funded by the priority program. The paper presents field observations and modelling efforts, extending from the stratosphere to the deep ocean. The research spans a large range of temporal and spatial scales, including the interaction of both climate components. In particular, radiative processes, the interaction of the changing ozone layer with large-scale atmospheric circulations, and changes in the sea ice cover are discussed. Climate and weather forecast models provide an insight into the water cycle and the climate change signals associated with synoptic cyclones. Investigations of the atmospheric boundary layer focus on the interaction between atmosphere, sea ice and ocean in the vicinity of polynyas and leads. The chapters dedicated to polar oceanography review the interaction between the ocean and ice shelves with regard to the freshwater input and discuss the changes in water mass characteristics, ventilation and formation rates, crucial for the deepest limb of the global, climate-relevant meridional overturning circulation. They also highlight the associated storage of anthropogenic carbon as well as the cycling of carbon, nutrients and trace metals in the ocean with special emphasis on the Weddell Sea.

First Evaluation of the Role of Salp Fecal Pellets on Iron Biogeochemistry

Planktonic grazers such as salps may have a dominant role in iron (Fe) cycling in surface waters of the Southern Ocean (SO). Salps have high ingestion rates and egest large, fast sinking fecal pellets (FPs) that potentially contribute to the vertical flux of carbon. In this study, we determined the impact of FPs from Salpa thompsoni, the most abundant salp in the SO, on Fe biogeochemistry. During the Polarstern expedition ANT-XXVII/3, salps were sampled from a large diatom bloom area in the Atlantic sector of the SO. Extensive work on carbon export and salp FPs export at the sampling location had shown that salps were a minor component of zooplankton and were responsible for only a 0.2% consumption of the daily primary production. Furthermore, at 100 m, export efficiency of salp FPs was ~2-3 fold higher than that of the bulk of sinking particulate organic carbon (POC). After collection, salps were maintained in 200 µm screened seawater and their FPs were collected for further experiments. To investigate whether the FPs release Fe and/or Fe-binding ligands into the filtered seawater (FSW) under different experimental conditions, they were either incubated in the dark or under full sunlight at in situ temperatures for 24h, or placed into the dark after a freeze/thaw treatment. We observed that none of the treatments caused release of dissolved Fe (dFe) or strong Fe ligands from the salp FPs. However, humic-substance like (HS-like) compounds, weak Fe ligands, were released at a rate of 8.2 ± 4.7 µg HS-like FP-1 d-1. Although the Fe content per salp FP was high at 0.33 ± 0.02 nmol dFe FP-1, the small contribution of salps to the zooplankton pool resulted in an estimated dFe export flux of 11.3 nmol Fe m-2 d-1 at 300 m. Since salp FPs showed an export efficiency at 100 m well above that shown by the bulk of sinking POC, our results suggest that in those areas of the SO where salps play a major role in the grazing of primary production, they could be actively contributing to the depletion of the dFe pool in surface water.