Characterization of the molecular mechanisms of silicon uptake in coccolithophores.

Coccolithophores are an important group of calcifying marine phytoplankton. Although coccolithophores are not silicified, some species exhibit a requirement for Si in the calcification process. These species also possess a novel protein (SITL) that resembles the SIT family of Si transporters found in diatoms. However, the nature of Si transport in coccolithophores is not yet known, making it difficult to determine the wider role of Si in coccolithophore biology. Here, we show that coccolithophore SITLs act as Na+ -coupled Si transporters when expressed in heterologous systems and exhibit similar characteristics to diatom SITs. We find that CbSITL from Coccolithus braarudii is transcriptionally regulated by Si availability and is expressed in environmental coccolithophore populations. However, the Si requirement of C. braarudii and other coccolithophores is very low, with transport rates of exogenous Si below the level of detection in sensitive assays of Si transport. As coccoliths contain only low levels of Si, we propose that Si acts to support the calcification process, rather than forming a structural component of the coccolith itself. Si is therefore acting as a micronutrient in coccolithophores and natural populations are only likely to experience Si limitation in circumstances where dissolved silicon (DSi) is depleted to extreme levels.

X-ray tomographic data of planktonic foraminifera species Globigerina bulloides from the Eastern Tropical Atlantic across Termination II.

Increased planktonic foraminifera shell weights were recorded during the course of Termination II at a tropical site off the shore of the Mauritanian coast. In order to investigate these increased shell mass values, a series of physicochemical analyses were performed, including X-ray computed tomography (CT). The data are given here. Furthermore, the relevant CT setup, scanning, reconstruction, and visualization methods are explained and the acquired datasets are given, together with 3D volumes and models of the scanned specimens.

Influence of surface ocean density on planktonic foraminifera calcification.

This study provides evidence that ambient seawater density influences calcification and may account for the observed planktonic foraminifera shell mass increase during glacial times. Volumes of weighed fossil Globigerina bulloides shells were accurately determined using X-ray Computer Tomography and were combined with water density reconstructions from Mg/Ca and δ18O measurements to estimate the buoyancy force exerted on each shell. After assessment of dissolution effects, the resulting relationship between shell mass and buoyancy suggests that heavier shells would need to be precipitated in glacial climates in order for these organisms to remain at their optimum living depth, and counterbalance the increased buoyant force of a denser, glacial ocean. Furthermore, the reanalysis of bibliographic data allowed the determination of a relationship between G. bulloides shell mass and ocean density, which introduces implications of a negative feedback mechanism for the uptake of atmospheric CO2 by the oceans.

Physiological responses of coccolithophores to abrupt exposure of naturally low pH deep seawater.

Upwelling is the process by which deep, cold, relatively high-CO2, nutrient-rich seawater rises to the sunlit surface of the ocean. This seasonal process has fueled geoengineering initiatives to fertilize the surface ocean with deep seawater to enhance productivity and thus promote the drawdown of CO2. Coccolithophores, which inhabit many upwelling regions naturally 'fertilized' by deep seawater, have been investigated in the laboratory in the context of ocean acidification to determine the extent to which nutrients and CO2 impact their physiology, but few data exist in the field except from mesocosms. Here, we used the Porcupine Abyssal Plain (north Atlantic Ocean) Observatory to retrieve seawater from depths with elevated CO2 and nutrients, mimicking geoengineering approaches. We tested the effects of abrupt natural deep seawater fertilization on the physiology and biogeochemistry of two strains of Emiliania huxleyi of known physiology. None of the strains tested underwent cell divisions when incubated in waters obtained from <1,000 m (pH = 7.99-8.08; CO2 = 373-485 p.p.m; 1.5-12 µM nitrate). However, growth was promoted in both strains when cells were incubated in seawater from ~1,000 m (pH = 7.9; CO2 ~560 p.p.m.; 14-17 µM nitrate) and ~4,800 m (pH = 7.9; CO2 ~600 p.p.m.; 21 µM nitrate). Emiliania huxleyi strain CCMP 88E showed no differences in growth rate or in cellular content or production rates of particulate organic (POC) and inorganic (PIC) carbon and cellular particulate organic nitrogen (PON) between treatments using water from 1,000 m and 4,800 m. However, despite the N:P ratio of seawater being comparable in water from ~1,000 and ~4,800 m, the PON production rates were three times lower in one incubation using water from ~1,000 m compared to values observed in water from ~4,800 m. Thus, the POC:PON ratios were threefold higher in cells that were incubated in ~1,000 m seawater. The heavily calcified strain NZEH exhibited lower growth rates and PIC production rates when incubated in water from ~4,800 m compared to ~1,000 m, while cellular PIC, POC and PON were higher in water from 4,800 m. Calcite Sr/Ca ratios increased with depth despite constant seawater Sr/Ca, indicating that upwelling changes coccolith geochemistry. Our study provides the first experimental and field trial of a geoengineering approach to test how deep seawater impacts coccolithophore physiological and biogeochemical properties. Given that coccolithophore growth was only stimulated using waters obtained from >1,000 m, artificial upwelling using shallower waters may not be a suitable approach for promoting carbon sequestration for some locations and assemblages, and should therefore be investigated on a site-by-site basis.

West Antarctic Ice Sheet retreat driven by Holocene warm water incursions.

Glaciological and oceanographic observations coupled with numerical models show that warm Circumpolar Deep Water (CDW) incursions onto the West Antarctic continental shelf cause melting of the undersides of floating ice shelves. Because these ice shelves buttress glaciers feeding into them, their ocean-induced thinning is driving Antarctic ice-sheet retreat today. Here we present a multi-proxy data based reconstruction of variability in CDW inflow to the Amundsen Sea sector, the most vulnerable part of the West Antarctic Ice Sheet, during the Holocene epoch (from 11.7 thousand years ago to the present). The chemical compositions of foraminifer shells and benthic foraminifer assemblages in marine sediments indicate that enhanced CDW upwelling, controlled by the latitudinal position of the Southern Hemisphere westerly winds, forced deglaciation of this sector from at least 10,400 years ago until 7,500 years ago-when an ice-shelf collapse may have caused rapid ice-sheet thinning further upstream-and since the 1940s. These results increase confidence in the predictive capability of current ice-sheet models.

Pliocene-Pleistocene evolution of sea surface and intermediate water temperatures from the southwest Pacific.

Over the last 5 million years, the global climate system has evolved toward a colder mean state, marked by large-amplitude oscillations in continental ice volume. Equatorward expansion of polar waters and strengthening temperature gradients have been detected. However, the response of the mid latitudes and high latitudes of the Southern Hemisphere is not well documented, despite the potential importance for climate feedbacks including sea ice distribution and low-high latitude heat transport. Here we reconstruct the Pliocene-Pleistocene history of both sea surface and Antarctic Intermediate Water (AAIW) temperatures on orbital time scales from Deep Sea Drilling Project Site 593 in the Tasman Sea, southwest Pacific. We confirm overall Pliocene-Pleistocene cooling trends in both the surface ocean and AAIW, although the patterns are complex. The Pliocene is warmer than modern, but our data suggest an equatorward displacement of the subtropical front relative to present and a poleward displacement of the subantarctic front of the Antarctic Circumpolar Current (ACC). Two main intervals of cooling, from ~3 Ma and ~1.5 Ma, are coeval with cooling and ice sheet expansion noted elsewhere and suggest that equatorward expansion of polar water masses also characterized the southwest Pacific through the Pliocene-Pleistocene. However, the observed trends in sea surface temperature and AAIW temperature are not identical despite an underlying link to the ACC, and intervals of unusual surface ocean warmth (~2 Ma) and large-amplitude variability in AAIW temperatures (from ~1 Ma) highlight complex interactions between equatorward displacements of fronts associated with the ACC and/or varying poleward heat transport from the subtropics.

Responses of the Emiliania huxleyi proteome to ocean acidification.

Ocean acidification due to rising atmospheric CO2 is expected to affect the physiology of important calcifying marine organisms, but the nature and magnitude of change is yet to be established. In coccolithophores, different species and strains display varying calcification responses to ocean acidification, but the underlying biochemical properties remain unknown. We employed an approach combining tandem mass-spectrometry with isobaric tagging (iTRAQ) and multiple database searching to identify proteins that were differentially expressed in cells of the marine coccolithophore species Emiliania huxleyi (strain NZEH) between two CO2 conditions: 395 (∼current day) and ∼1340 p.p.m.v. CO2. Cells exposed to the higher CO2 condition contained more cellular particulate inorganic carbon (CaCO3) and particulate organic nitrogen and carbon than those maintained in present-day conditions. These results are linked with the observation that cells grew slower under elevated CO2, indicating cell cycle disruption. Under high CO2 conditions, coccospheres were larger and cells possessed bigger coccoliths that did not show any signs of malformation compared to those from cells grown under present-day CO2 levels. No differences in calcification rate, particulate organic carbon production or cellular organic carbon: nitrogen ratios were observed. Results were not related to nutrient limitation or acclimation status of cells. At least 46 homologous protein groups from a variety of functional processes were quantified in these experiments, of which four (histones H2A, H3, H4 and a chloroplastic 30S ribosomal protein S7) showed down-regulation in all replicates exposed to high CO2, perhaps reflecting the decrease in growth rate. We present evidence of cellular stress responses but proteins associated with many key metabolic processes remained unaltered. Our results therefore suggest that this E. huxleyi strain possesses some acclimation mechanisms to tolerate future CO2 scenarios, although the observed decline in growth rate may be an overriding factor affecting the success of this ecotype in future oceans.

Effect of carbonate chemistry alteration on the early embryonic development of the Pacific oyster (Crassostrea gigas).

Ocean acidification, due to anthropogenic CO2 absorption by the ocean, may have profound impacts on marine biota. Calcareous organisms are expected to be particularly sensitive due to the decreasing availability of carbonate ions driven by decreasing pH levels. Recently, some studies focused on the early life stages of mollusks that are supposedly more sensitive to environmental disturbances than adult stages. Although these studies have shown decreased growth rates and increased proportions of abnormal development under low pH conditions, they did not allow attribution to pH induced changes in physiology or changes due to a decrease in aragonite saturation state. This study aims to assess the impact of several carbonate-system perturbations on the growth of Pacific oyster (Crassostrea gigas) larvae during the first 3 days of development (until shelled D-veliger larvae). Seawater with five different chemistries was obtained by separately manipulating pH, total alkalinity and aragonite saturation state (calcium addition). Results showed that the developmental success and growth rates were not directly affected by changes in pH or aragonite saturation state but were highly correlated with the availability of carbonate ions. In contrast to previous studies, both developmental success into viable D-shaped larvae and growth rates were not significantly altered as long as carbonate ion concentrations were above aragonite saturation levels, but they strongly decreased below saturation levels. These results suggest that the mechanisms used by these organisms to regulate calcification rates are not efficient enough to compensate for the low availability of carbonate ions under corrosive conditions.