Seawater carbonate chemistry and calcification of two mediterranean cold-water coral species in a laboratory experiment

Deep-water ecosystems are characterized by relatively low carbonate concentration values and, due to ocean acidification (OA), these habitats might be among the first to be exposed to undersaturated conditions in the forthcoming years. However, until now, very few studies have been conducted to test how cold-water coral (CWC) species react to such changes in the seawater chemistry. The present work aims to investigate the mid-term effect of decreased pH on calcification of the two branching CWC species most widely distributed in the Mediterranean, Lophelia pertusa and Madrepora oculata. No significant effects were observed in the skeletal growth rate, microdensity and porosity of both species after 6 months of exposure. However, while the calcification rate of M. oculata was similar for all colony fragments, a heterogeneous skeletal growth pattern was observed in L. pertusa, the younger nubbins showing higher growth rates than the older ones. A higher energy demand is expected in these young, fast-growing fragments and, therefore, a reduction in calcification might be noticed earlier during long-term exposure to acidified conditions.

Abundance and biomass of protozooplankton from the iron-induced phytoplankton bloom during the EisenEx experiment in 2000 in the Southern Ocean

The responses of larger (>50 µm in diameter) protozooplankton groups to a phytoplankton bloom induced by in situ iron fertilization (EisenEx) in the Polar Frontal Zone (PFZ) of the Southern Ocean in austral spring are presented. During the 21 days of the experiment, samples were collected from seven discrete depths in the upper 150 m inside and outside the fertilized patch for the enumeration of acantharia, foraminifera, radiolaria, heliozoa, tintinnid ciliates and aplastidic thecate dinoflagellates. Inside the patch, acantharian numbers increased twofold, but only negligibly in surrounding waters. This finding is of major interest, since acantharia are suggested to be involved in the formation of barite (BaSO_4 ) found in sediments and which is a palaeoindicator of both ancient and modern high productivity regimes. Foraminifera increased significantly in abundance inside and outside the fertilized patch. However the marked increase of juveniles after a full moon event suggests a lunar periodicity in the reproduction cycle of some foraminiferan species rather than a reproductive response to enhanced food availability. In contrast, adult radiolaria showed no clear trend during the experiment, but juveniles increased threefold indicating elevated reproduction. Aplastidic thecate dinoflagellates almost doubled in numbers and biomass, but also increased outside the patch. Tintinnid numbers decreased twofold, although biomass remained constant due to a shift in the size spectrum. Empty tintinnid loricae, however, increased by a factor of two indicating that grazing pressure on this group mainly by copepods intensified during EisenEx. The results show that iron-fertilization experiments can shed light on the biology and the role of these larger protists in pelagic ecosystem which will improve their use as proxies in palaeoceanography.

Gasterosteus aculeatus egg traits and fish lengths vs. experiment setup

1) Our study addresses the role of non-genetic and genetic inheritance in shaping the adaptive potential of populations under a warming ocean scenario. We used a combined experimental approach (transgenerational plasticity and quantitative genetics) to partition the relative contribution of maternal vs. paternal (additive genetic) effects to offspring body size (a key component of fitness), and investigated a potential physiological mechanism (mitochondrial respiration capacities) underlying whole organism growth/size responses. 2) In very early stages of growth (up to 30 days), offspring body size of marine sticklebacks benefited from maternal transgenerational plasticity (TGP): offspring of mothers acclimated to17°C were larger when reared at 17°C, and offspring of mothers acclimated to 21°C were larger when reared at 21°C. The benefits of maternal TGP on body size were stronger and persisted longer (up to 60 days) for offspring reared in the warmer (21°C) environment, suggesting that maternal effects will be highly relevant for climate change scenarios in this system. 3) Mitochondrial respiration capacities measured on mature offspring (F1 adults) matched the pattern of TGP for juvenile body size, providing an intuitive mechanistic basis for the maternal acclimation persisting into adulthood. Size differences between temperatures seen at early growth stages remained in the F1 adults, linking offspring body size to maternal inheritance of mitochondria. 4) Lower maternal variance components in the warmer environment were mostly driven by mothers acclimated to ambient (colder) conditions, further supporting our tenet that maternal effects were stronger at elevated temperature. Importantly, all parent-offspring temperature combination groups showed genotype x environment (GxE) interactions, suggesting that reaction norms have the potential to evolve. 5) To summarise, transgenerational plasticity and genotype x environment interactions work in concert to mediate impacts of ocean warming on metabolic capacity and early growth of marine sticklebacks. TGP can buffer short-term detrimental effects of climate warming and may buy time for genetic adaptation to catch up, therefore markedly contributing to the evolutionary potential and persistence of populations under climate change.

A 34-month sediment trap experiment at the Cape Verde Ocean Observatory (CVOO, December 2009 - October 2012)

Anticyclonic mesoscale eddies (ACME) have been proposed as a mechanism by which new nutrients are episodically delivered into the euphotic zone, thereby enhancing new production as well as shifting phytoplankton community structure. In this paper, we report on a 34-month sediment trap experiment at the Cape Verde Ocean Observatory (CVOO; ca. 18°N, 24°E; December 2009-October 2012), occasionally influenced by ACME passages. The typically oligotrophic, weakly seasonal particle flux pattern at the CVOO is strongly modified by the appearance of a highly productive and low oxygen ACME. Out of four recorded diatom flux maxima at CVOO, three were associated with the passage of ACMEs. The recorded diatom maxima events support the view that local ACME dynamics promotes upward nutrient supply into the euphotic zone leading to a rapid response of diatoms. This response is clearly reflected by the flux seasonality: between 40% and 60% of the total annual diatom flux at the CVOO site was intercepted in a relatively short time interval (<60 days). A highly diverse diatom community characterized the diatom fluxes throughout. Along with the ACME passages, small species of the genus Nitzschia, and Thalassionema nitzschioides var. parva dominated and delivered a major portion of the opal and organic carbon into deeper waters at site CVOO. Several pelagic, warm-water background species became dominant during intervals with low nutrient availability in the euphotic zone. Results of our interannual time-series suggest that ACMEs impact on total diatom production and the species-specific composition of the assemblage north of the Cave Verde Islands, and can strengthen the biological pump in open-ocean, oligotrophic subtropical regions of the world ocean. Our observations are useful for testing biogeochemical ocean models and will also help in improving the knowledge of processes and mechanisms behind interannual time-series of bulk components and microorganisms in pelagic and hemipelagic ocean areas.

Dimethylsulphide (DMS), dimethylsulphonioproionate (DMSP) and halocarbon concentrations taken from 25m integrated samples during the 2011 SOPRAN Bergen Raunefjord Mesocosm experiment, Norway

The human-induced rise in atmospheric carbon dioxide since the industrial revolution has led to increasing oceanic carbon uptake and changes in seawater carbonate chemistry, resulting in lowering of surface water pH. In this study we investigated the effect of increasing CO2 partial pressure (pCO2) on concentrations of volatile biogenic dimethylsulfide (DMS) and its precursor dimethylsulfoniopropionate (DMSP), through monoculture studies and community pCO2 perturbation. DMS is a climatically important gas produced by many marine algae: it transfers sulfur into the atmosphere and is a major influence on biogeochemical climate regulation through breakdown to sulfate and formation of subsequent cloud condensation nuclei (CCN). Overall, production of DMS and DMSP by the coccolithophore Emiliania huxleyi strain RCC1229 was unaffected by growth at 900 µatm pCO2, but DMSP production normalised to cell volume was 12 % lower at the higher pCO2 treatment. These cultures were compared with community DMS and DMSP production during an elevated pCO2 mesocosm experiment with the aim of studying E. huxleyi in the natural environment. Results contrasted with the culture experiments and showed reductions in community DMS and DMSP concentrations of up to 60 and 32 % respectively at pCO2 up to 3000 µatm, with changes attributed to poorer growth of DMSP-producing nanophytoplankton species, including E. huxleyi, and potentially increased microbial consumption of DMS and dissolved DMSP at higher pCO2. DMS and DMSP production differences between culture and community likely arise from pH affecting the inter-species responses between microbial producers and consumers.

Oceanographic investiations from the Sea of Ochotsk

The Sea of Okhotsk is a marginal sea of the Pacific Ocean, which is characterized by strong variations in the productivity and sediment supply due to sea ice transport and river input. Furthermore the variations in the hydrological cycle determine the formation of the SOIW (Sea of Okhotsk Intermediate Water) which plays an important role in the ventilation processes in the intermediate water of the N-Pacific. Isotope data measured on planktonic and benthic foraminifera, sedimentological and geochemical studies of sediment cores and surface samples from the Sea of Okhotsk are used to reconstruct the paleoceanography during the past 350.000 years. The dating and correlation of the sediments are based on oxygen isotope stratigraphy, absolute ages, magnetic susceptibility as well as a detailled tephrachronology of the entire basin. The sedimentation rates are characterized by temporal and spatial variations. The maximum sedimentation rate takes place at the continental slope off Sakhalin due to the input of the Amur River, the sea ice drift and the high productivity. The sedimentation rate in the eastern part of the Sea of Okhotsk is generelly high because of the influence of the nutrient-rich Kamchatka Current. In the central and northern parts of the Sea of Okhotsk, areas with low productivity and reduced terrestrial supply, the sedimentation rate is the lowest. The analyses of the surface sediment samples make it possible to characterize the (sub)- recent sediment supply and transportation processes. The bulk sediment measurements, isotope data and the accumulation rate of ice-rafted debris (IRD) show a dominant sea ice cover and a region with a high productivity as well as a high Amur River input in the western part of the sea. The eastern part of the Sea of Okhotsk, however, is marked by the predominance of warm and nutrient-rich water masses coming from the Kamchatka Current which restricts the sea ice cover. This is reflected in low content of ice-rafted debris and high productivity proxies as well as in isotope data. The deposits of the Sea of Okhotsk are characterized by terrestrial, biogenic and volcanogenic sediment input which varies temporally and spatially. Here, the sedimentation pattern is dominated by the terrestrial input. Bulk sediment measurements and sample analyses of the > 63 micron particle input make it possible to distinguish glacial and interglacial fluctuations. The sedimentation processes during glacial times are determined by a high content of ice-rafted debris, whereas the primary production is higher during interglacial periods. During the last glacial/interglacial cycle the IRD-distribution pattern indicates a strong sea ice transport in the western part and in large areas of the open sea in the eastern part of the Sea of Okhotsk with a relatively constant ice-drift system. The IRD flux in sediments of the oxygen isotope Stage 6 reflects a new sedimentation pattern in the eastern part of the sea. This high IRD accumulation rate indicates ice advances beyond the shelf margin and an iceberg transport from NE-E direction into the Sea of Okhotsk. The several large, brief, negative anomalies in d13C values of Neogloboquadrina pachyderma (s) show releases of methane from basin sediments which correspond to periods of relative sea level falls. The high sedimentation rates on the Sakhalin slope allow insights into the climatic history in Holocene and indicate shorter-scale variations oscillation in Stage 3, which correlate with the global climatic changes. These variations are described as Dansgaard-Oeschger cycles in Greenland ice cores and as Heinrich-Events in several marine sediment cores from the N-Atlantic.

Experiment: Acidified seawater impacts sea urchin larvae pH regulatory systems relevant for calcification

Calcifying echinoid larvae respond to changes in seawater carbonate chemistry with reduced growth and developmental delay. To date, no information exists on how ocean acidification acts on pH homeostasis in echinoderm larvae. Understanding acid-base regulatory capacities is important because intracellular formation and maintenance of the calcium carbonate skeleton is dependent on pH homeostasis. Using H(+)-selective microelectrodes and the pH-sensitive fluorescent dye BCECF, we conducted in vivo measurements of extracellular and intracellular pH (pH(e) and pH(i)) in echinoderm larvae. We exposed pluteus larvae to a range of seawater CO(2) conditions and demonstrated that the extracellular compartment surrounding the calcifying primary mesenchyme cells (PMCs) conforms to the surrounding seawater with respect to pH during exposure to elevated seawater pCO(2). Using FITC dextran conjugates, we demonstrate that sea urchin larvae have a leaky integument. PMCs and spicules are therefore directly exposed to strong changes in pH(e) whenever seawater pH changes. However, measurements of pH(i) demonstrated that PMCs are able to fully compensate an induced intracellular acidosis. This was highly dependent on Na(+) and HCO(3)(-), suggesting a bicarbonate buffer mechanism involving secondary active Na(+)-dependent membrane transport proteins. We suggest that, under ocean acidification, maintained pH(i) enables calcification to proceed despite decreased pH(e). However, this probably causes enhanced costs. Increased costs for calcification or cellular homeostasis can be one of the main factors leading to modifications in energy partitioning, which then impacts growth and, ultimately, results in increased mortality of echinoid larvae during the pelagic life stage.

Seawater carbonate chemistry and Strongylocentrotus purpuratus body length and gene expression pattern changes during experiments, 2011

Extensive use of fossil fuels is leading to increasing CO2 concentrations in the atmosphere and causes changes in the carbonate chemistry of the oceans which represents a major sink for anthropogenic CO2. As a result, the oceans' surface pH is expected to decrease by ca. 0.4 units by the year 2100, a major change with potentially negative consequences for some marine species. Because of their carbonate skeleton, sea urchins and their larval stages are regarded as likely to be one of the more sensitive taxa. In order to investigate sensitivity of pre-feeding (2 days post-fertilization) and feeding (4 and 7 days post-fertilization) pluteus larvae, we raised Strongylocentrotus purpuratus embryos in control (pH 8.1 and pCO2 41 Pa e.g. 399 µatm) and CO2 acidified seawater with pH of 7.7 (pCO2 134 Pa e.g. 1318 µatm) and investigated growth, calcification and survival. At three time points (day 2, day 4 and day 7 post-fertilization), we measured the expression of 26 representative genes important for metabolism, calcification and ion regulation using RT-qPCR. After one week of development, we observed a significant difference in growth. Maximum differences in size were detected at day 4 (ca. 10 % reduction in body length). A comparison of gene expression patterns using PCA and ANOSIM clearly distinguished between the different age groups (Two way ANOSIM: Global R = 1) while acidification effects were less pronounced (Global R = 0.518). Significant differences in gene expression patterns (ANOSIM R = 0.938, SIMPER: 4.3% difference) were also detected at day 4 leading to the hypothesis that differences between CO2 treatments could reflect patterns of expression seen in control experiments of a younger larva and thus a developmental artifact rather than a direct CO2 effect. We found an up regulation of metabolic genes (between 10 to 20% in ATP-synthase, citrate synthase, pyruvate kinase and thiolase at day 4) and down regulation of calcification related genes (between 23 and 36% in msp130, SM30B, SM50 at day 4). Ion regulation was mainly impacted by up regulation of Na+/K+-ATPase at day 4 (15%) and down regulation of NHE3 at day 4 (45%). We conclude that in studies in which a stressor induces an alteration in the speed of development, it is crucial to employ experimental designs with a high time resolution in order to correct for developmental artifacts. This helps prevent misinterpretation of stressor effects on organism physiology.