Nitrification rates in the NW Mediterranean Sea

During spring, ammonium oxidation and nitrite oxidation rates were measured in the NW basin of the Mediterranean Sea, from mesotrophic sites (Ligurian Sea and Gulf of Lions) to oligotrophic sites (Balearic Islands). Nitrification rates (average values for 37 measurements) ranged from 72 to 144 nmol of N oxidised/l/d, except in the Rhône River plume area where the rates increased to 264-504 nmol/l/d because of the riverine inputs of nitrogen. Maximal rates were located around the peak of nitrite within the nitracline at about 40 to 60 m and just above the phosphacline. At 1 station, relatively high values of nitrification (50 to 130 nmol/l/d) were also measured deep in the water column (240 m). Day-to-day variations were measured demonstrating the response within a few hours to hydrological stress (wind-induced mixing of the water column) and showing the role of hydrological characteristics on the distribution of nitrification rates. Because of the homogenous temperature (13°C) in the Mediterranean Sea, the spatial (geographical and vertical) fluctuations of nitrifying rates were linked to the presence of substrate due to mineralisation processes and/or Rhône River inputs. We estimate the contribution of nitrate produced by nitrification to the N demand of phytoplankton to range from 16% at mesotrophic to 61% at oligotrophic stations.

Abundance and biomass of phytoplankton in the Black Sea near Gelendzhik in 2002-2005

It is shown that in 2002-2005 mass development of coccolithofore Emiliania huxleyi on the Gelendzhik shelf (northeast Black Sea) occurred annually and in May-June its abundance reached 1500000 cells/l. In 2004-2005 bloom of E. huxleyi was accompanied by mass development of diatom alga Chaetoceros subtilis var. abnormis f. simplex (600000-900000 cells/l). For the first time it was registered as a dominating form of Black Sea phytoplankton. Small flagellates and picoplankton algae played a noticeable role in phytoplankton throughout the entire period of the studies. Meanwhile in the early summer period the bulk of biomass consisted of coccolithophores (50-60%), while in the late summer period diatomaceous algae dominated (50-70%). Among ecological factors that favor coccolithophore development one may note microstratification of the upper mixed layer at a high illumination level and high temperature in surface waters (18-21°C). Terrigenous runoff during the rainy period had a negative effect on E. huxleyi development, while storms dispersed the population over the upper mixed layer. A wind-induced near-shore upwelling stimulated development of diatoms.

Age determination and paleotemperatures of sediment cores TTR17-1_384B and TTR17-1_436B in the Alboran Sea

Climate conditions in the westernmost Mediterranean (Alboran Sea basin) over the last two millennia have been reconstructed through integration of molecular proxies applied for the first time in this region at such high resolution. Two temperature proxies, one based on isoprenoid membrane lipids of marine Thaumarchaeota (TEXH86-tetraether index of compounds consisting of 86 carbons) and the other on alkenones produced by haptophytes (UK'37 ratio) were applied to reconstruct sea surface temperature (SST). Both records reveal a progressive long term decline in SST over the last two millennia and an increased rate of warming during the second half of the twentieth century. This is in accord with previous temperature reconstructions for the Northern Hemisphere. TEXH86 temperature values are higher than those inferred from UK'37, probably due to differences in the bloom season of haptophytes and Thaumarchaeota, and reflect summer SST. The branched vs. isoprenoid tetraether index (BIT index) suggests a low contribution of soil organic matter (OM) to the sedimentary OM. The stable carbon isotopic composition of long chain n-alkanes indicates a predominant C3 plant contribution, with no major change in vegetation over the last 2000 yr. The distribution of long chain 1,14-diols (most likely sourced by Proboscia species in this setting) provided insight into variation in upwelling conditions during the last 2000 yr and depicts a correlation with the North Atlantic Oscillation (NAO) index, providing evidence of enhanced wind induced upwelling during periods of a persistent positive mode of the NAO.

Figure 2. Downcore variations of total diatom concentration in core MD02-2529

The modern eastern equatorial Pacific (EEP) is a major natural source for atmospheric carbon dioxide and is thought to be connected to high-latitude ocean dynamics by oceanic teleconnections on glacial-interglacial timescales. A wealth of sedimentary records aiming at reconstructing last Quaternary changes in primary productivity and nutrient utilization have been devoted to understanding those linkages between the EEP and other distant oceanic areas. Most of these records are, however, clustered in the pelagic EEP cold tongue, with comparatively little attention devoted to coastal areas. Here we present downcore measurements of the composition and concentration of the diatom assemblage together with opal (biogenic silica) concentration at site MD02-2529 recovered in the coastal Panama Basin. Piston core MD02-2529, collected in an area affected by a multitude of processes, provides evidence for strong variations in diatom production at the millennial timescale during the last glacial cycle. The maxima in total diatom concentration occurred during the early marine isotopic stage (MIS) 4 as well as during the MIS 4/3 transition and MIS 3. Rapid changes in diatom concentrations during the MIS 3 mimics Bond cycles as independently recorded by the SSS estimation derived from planktonic foraminifera from the same core. Such patterns indicate a clear linkage between diatom production in the coastal EEP and rapid climate changes in the high-latitude North Atlantic. In parallel, the long-term succession of the diatom community from coastal diatoms, predominantly thriving during MIS 5 and 4, towards pelagic diatoms, dominant during MIS 3 and 2, points to a long-term change in the surface hydrology. During Heinrich Events, diatoms strongly reduced their production, probably due to enhanced stratification in the upper water column. After the last glacial maximum, diatom production and valve preservation strongly decreased in response to the advection of nutrient (H2SiO4)-depleted, warmer water masses. Our high-resolution record highlights how regional climatic processes can modulate rapid changes in siliceous primary production as triggered by wind-induced local upwelling, indicating that millennial climatic variability can overtake other prominent hydrological processes such as those related to silicic acid leakage.

Late Eocene to Early Oligocene magnetostratigraphic chron boundaries of ODP Hole 119-744A (Table 1)

The earliest Oligocene (~33.5 Ma) is marked by a major step in the long-term transition from an ice-free to glaciated world. The transition, characterized by both cooling and ice-sheet growth, triggered a transient but extreme glacial period designated Oi-1. High-resolution isotope records suggest that Oi-1 lasted for roughly 400,000 yr (the duration of magnetochron 13N) before partially abating, and that it was accompanied by an ocean-wide carbon isotope anomaly of 0.75?. One hypothesis relates the carbon isotope anomaly to enhanced export production brought about by climate-induced intensification of wind stress and upwelling, particularly in the Southern Ocean. To understand how this climatic event affected export production in the Southern Ocean, biogenic silica (opal) and carbonate accumulation rates were computed for the sub-polar Indian Ocean using deep-sea cores from ODP Site 744, Kerguelen Plateau. Our findings suggest that net productivity in this region increased by several fold in response to the Oi-1 glaciation. In addition, calcareous primary producers dominant in the Late Eocene were partially replaced by opaline organisms suggesting a trend toward seasonally greater surface divergence and upwelling in this sector of the Southern Ocean. We attribute these changes to intensification of atmospheric=oceanic circulation brought about by high-latitude cooling and the appearance of a full-scale continental ice-sheet on East Antarctica. Higher terrigenous sediment accumulation rates support the idea that wind-induced changes in regional productivity were augmented by an increased supply of glacial dust and debris that provided limiting micro-nutrients (e.g., iron-rich dust particles). We speculate that the rapid changes in biogenic sediment accumulation in the Southern Ocean and other upwelling-dominated regions contributed to the ocean-wide positive carbon isotope anomaly by temporarily increasing the burial rate of organic carbon relative to carbonate carbon. The changes in burial rates, in turn, may have produced a positive feedback on climate by briefly drawing down atmospheric pCO2 .

(Table 2) Age determination of sediment core NIOP-C2_929

In this study we present a sea surface temperature (SST) record from the western Arabian Sea for the last 20,000 years. We produced centennial-scale d18O and Mg/Ca SST time series of core NIOP929 with focus on the glacial-interglacial transition. The western Arabian Sea is influenced by the seasonal NE and SW monsoon wind systems. Lowest SSTs occur during the SW monsoon season because of upwelling of cold water, and highest SSTs can be found in the low-productivity intermonsoon season. The Mg/Ca-based temperature record reflects the integrated SST of the SW and NE monsoon seasons. The results show a glacial-interglacial SST difference of ~2°C, which is corroborated by findings from other Arabian Sea cores. At 19 ka B.P. a yet undescribed warm event of several hundred years duration is found, which is also reflected in the d18O record. A second centennial-scale high SST/low d18O event is observed at 17 ka B.P. This event forms the onset of the stepwise yet persistent trend toward Holocene temperatures. Highest Mg/Ca-derived SSTs in the NIOP929 record occurred between 13 and 10 ka B.P. Interglacial SST is ~24°C, indicating influence of upwelling. The onset of Arabian Sea warming occurs when the North Atlantic is experiencing minimum temperatures. The rapid temperature variations at 19, 17, and 13 ka B.P. are difficult to explain with monsoon changes alone and are most likely also linked to regional hydrographic changes, such as trade wind induced variations in warm water advection.

Georeferenced standardized apparent wave power along the coastline of Cabrera National Park for analysis of essential habitats of littoral species

Wind- induced exposure is one of the major forces shaping the geomorphology and biota in coastal areas. The effect of wave exposure on littoral biota is well known in marine environments (Ekebon et al., 2003; Burrows et al., 2008). In the Cabrera Archipelago National Park wave exposure has demostrated to have an effect on the spatial distribution of different stages of E.marginatus (Alvarez et al., 2010). Standarized average wave exposures during 2008 along the Cabrera Archipelago National park coast line were calculated to be applied in studies of littoral species distribution within the archipelago. Average wave exposure (or apparent wave power) was calculated for points located 50 m equidistant on the coastline following the EXA methodology (EXposure estimates for fragmented Archipelagos) (Ekebon et al., 2003). The average wave exposures were standardized from 1 to 100 (minimum and maximum in the area), showing coastal areas with different levels of mea wave exposure during the year. Input wind data (direction and intensity) from 2008 was registered at the Cabrera mooring located north of Cabrera Archipelago. Data were provided by IMEDEA (CSIC-UIB, TMMOS http://www.imedea.uib-csic.es/tmoos/boyas/). This cartography has been developed under the framework of the project EPIMHAR, funded by the National Park's Network (Spanish Ministry of Environment, Maritime and Rural Affairs, reference: 012/2007 ). Part of this work has been developed under the research programs funded by "Fons de Garantia Agrària i Pesquera de les Illes Balears (FOGAIBA)".

Two intertidal, non-calcifying macroalgae (Palmaria palmata and Saccharina latissima) show complex and variable responses to short-term CO2 acidification

Ocean acidification, the result of increased dissolution of carbon dioxide (CO2) in seawater, is a leading subject of current research. The effects of acidification on non-calcifying macroalgae are, however, still unclear. The current study reports two 1-month studies using two different macroalgae, the red alga Palmaria palmata (Rhodophyta) and the kelp Saccharina latissima (Phaeophyta), exposed to control (pHNBS = 8.04) and increased (pHNBS = 7.82) levels of CO2-induced seawater acidification. The impacts of both increased acidification and time of exposure on net primary production (NPP), respiration (R), dimethylsulphoniopropionate (DMSP) concentrations, and algal growth have been assessed. In P. palmata, although NPP significantly increased during the testing period, it significantly decreased with acidification, whereas R showed a significant decrease with acidification only. S. latissima significantly increased NPP with acidification but not with time, and significantly increased R with both acidification and time, suggesting a concomitant increase in gross primary production. The DMSP concentrations of both species remained unchanged by either acidification or through time during the experimental period. In contrast, algal growth differed markedly between the two experiments, in that P. palmata showed very little growth throughout the experiment, while S. latissima showed substantial growth during the course of the study, with the latter showing a significant difference between the acidified and control treatments. These two experiments suggest that the study species used here were resistant to a short-term exposure to ocean acidification, with some of the differences seen between species possibly linked to different nutrient concentrations between the experiments.

Modeled integrated responses of ppERK1 and ppERK2 of normal and elevated ERK1 and ERK2 levels (Figure 5b)

The present dataset contain source data for Figure 5b from Schilling et al., 2009. Cell fate decisions are regulated by the coordinated activation of signalling pathways such as the extracellular signal-regulated kinase (ERK) cascade, but contributions of individual kinase isoforms are mostly unknown. The authors combined quantitative data from erythropoietin-induced pathway activation in primary erythroid progenitor (colony-forming unit erythroid stage, CFU-E) cells with mathematical modelling, in order to predict and experimentally confirmed a distributive ERK phosphorylation mechanism in CFU-E cells. The authors found evidences that double-phosphorylated ERK1 attenuates proliferation beyond a certain activation level, whereas activated ERK2 enhances proliferation with saturation kinetics. They show integrated responses of double-phosphorylated ERK1 and ERK2 that were calculated for different Epo concentrations for the original model as well as for models with elevated ERK1 or ERK2 levels.

Area-averaged mean seasonal cycles of chlorophyll-a concentration, sea surface temperature, alongshore wind stress, westward wind speed and dust component of the aerosol optical depth at 550 nm

Nutrient supply in the area off Northwest Africa is mainly regulated by two processes, coastal upwelling and deposition of Saharan dust. In the present study, both processes were analyzed and evaluated by different methods, including cross-correlation, multiple correlation, and event statistics, using remotely sensed proxies of the period from 2000 to 2008 to investigate their influence on the marine environment. The remotely sensed chlorophyll-a concentration was used as a proxy for the phytoplankton biomass stimulated by nutrient supply into the euphotic zone from deeper water layers and from the atmosphere. Satellite-derived alongshore wind stress and sea-surface temperature were applied as proxies for the strength and reflection of coastal upwelling processes. The westward wind and the dust component of the aerosol optical depth describe the transport direction of atmospheric dust and the atmospheric dust column load. Alongshore wind stress and induced upwelling processes were most significantly responsible for the surface chlorophyll-a variability, accounting for about 24% of the total variance, mainly in the winter and spring due to the strong north-easterly trade winds. The remotely sensed proxies allowed determination of time lags between biological response and its forcing processes. A delay of up to 16 days in the surface chlorophyll-a concentration due to the alongshore wind stress was determined in the northern winter and spring. Although input of atmospheric iron by dust storms can stimulate new phytoplankton production in the study area, only 5% of the surface chlorophyll-a variability could be ascribed to the dust component in the aerosol optical depth. All strong desert storms were identified by an event statistics in the time period from 2000 to 2008. The 57 strong storms were studied in relation to their biological response. Six events were clearly detected in which an increase of chlorophyll-a was caused by Saharan dust input and not by coastal upwelling processes. Time lags of <8 days, 8 days, and 16 days were determined. An increase in surface chlorophyll-a concentration of up to 2.4 mg m**3 after dust storms in which the dust component of the aerosol optical depth was up to 0.9 was observed.

Glacial-interglacial bioproductivity of the Mexican Margin

Sedimentary accumulation of biogenic components (organic carbon, opal, and biogenic barium) on the northwestern Mexican margin declined during every glacial interval of the past 140 kyr, indicating decreases in upwelling-induced productivity during cold periods. The glacial-interglacial contrasts in upwelling on this margin are attributed to reversals in land-ocean thermal contrast, the waxing and waning of the Laurentide Ice Sheet, and consequent responses of the western hemisphere wind fields. This scenario is consistent with three independent lines of evidence: terrestrial paleoclimatic data, general circulation model results, and our marine records. This pattern of glacial-interglacial variability in upwelling off NW Mexico is opposite to that observed in other low-latitude and midlatitude upwelling areas, such as the eastern equatorial Pacific. These results add to a growing pool of observations that the response of oceanic upwelling to glacial climatic forcing has been regionally variable.

Seawater carbonate chemistry, gross photosynthesis and metabolically induced rate of pH change during experiments with macroalgae, 2012

Ocean acidification (OA) is a reduction in oceanic pH due to increased absorption of anthropogenically produced CO2. This change alters the seawater concentrations of inorganic carbon species that are utilized by macroalgae for photosynthesis and calcification: CO2 and HCO3 increase; CO32 decreases. Two common methods of experimentally reducing seawater pH differentially alter other aspects of carbonate chemistry: the addition of CO2 gas mimics changes predicted due to OA, while the addition of HCl results in a comparatively lower [HCO3]. We measured the short-term photosynthetic responses of five macroalgal species with various carbon-use strategies in one of three seawater pH treatments: pH 7.5 lowered by bubbling CO2 gas, pH 7.5 lowered by HCl, and ambient pH 7.9. There was no difference in photosynthetic rates between the CO2, HCl, or pH 7.9 treatments for any of the species examined. However, the ability of macroalgae to raise the pH of the surrounding seawater through carbon uptake was greatest in the pH 7.5 treatments. Modeling of pH change due to carbon assimilation indicated that macroalgal species that could utilize HCO3 increased their use of CO2 in the pH 7.5 treatments compared to pH 7.9 treatments. Species only capable of using CO2 did so exclusively in all treatments. Although CO2 is not likely to be limiting for photosynthesis for the macroalgal species examined, the diffusive uptake of CO2 is less energetically expensive than active HCO3 uptake, and so HCO3-using macroalgae may benefit in future seawater with elevated CO2.

The photo-physiological response of a model cnidarian-dinoflagellate symbiosis to CO2-induced acidification at the cellular level

We measured the relationship between CO2-induced seawater acidification, photo-physiological performance and intracellular pH (pHi) in a model cnidarian-dinoflagellate symbiosis - the sea anemone Aiptasia sp. -under ambient (289.94 ± 12.54 µatm), intermediate (687.40 ± 25.10 µatm) and high (1459.92 ± 65.51 µatm) CO2 conditions. These treatments represented current CO2 levels, in addition to CO2 stabilisation scenarios IV and VI provided by the Intergovernmental Panel on Climate Change (IPCC). Anemones were exposed to each treatment for two months and sampled at regular intervals. At each time-point we measured a series of physiological responses: maximum dark-adapted fluorescent yield of PSII (Fv/Fm), gross photosynthetic rate, respiration rate, symbiont population density, and light-adapted pHi of both the dinoflagellate symbiont and isolated host anemone cell. We observed increases in all but one photo-physiological parameter (Pgross:R ratio). At the cellular level, increases in light-adapted symbiont pHi were observed under both intermediate and high CO2 treatments, relative to control conditions (pHi 7.35 and 7.46 versus pHi 7.25, respectively). The response of light-adapted host pHi was more complex, however, with no change observed under the intermediate CO2 treatment, but a 0.3 pH-unit increase under the high CO2 treatment (pHi 7.19 and 7.48, respectively). This difference is likely a result of a disproportionate increase in photosynthesis relative to respiration at the higher CO2 concentration. Our results suggest that, rather than causing cellular acidosis, the addition of CO2 will enhance photosynthetic performance, enabling both the symbiont and host cell to withstand predicted ocean acidification scenarios.

Adaptive variability to low-pH river discharges in Acartia tonsa and stress responses to high PCO2 conditions

Environmental transitions leading to spatial physical-chemical gradients are of ecological and evolutionary interest because they are able to induce variations in phenotypic plasticity. Thus, the adaptive variability to low-pH river discharges may drive divergent stress responses [ingestion rates (IR) and expression of stress-related genes such as Heat shock protein 70 (Hsp70) and Ferritin] in the neritic copepod Acartia tonsa facing changes in the marine chemistry associated to ocean acidification (OA). These responses were tested in copepod populations inhabiting two environments with contrasting carbonate system parameters (an estuarine versus coastal area) in the Southern Pacific Ocean, and assessing an in situ and 96-h experimental incubation under conditions of high pressure of CO2 (PCO2 1200 ppm). Adaptive variability was a determining factor in driving variability of copepods' responses. Thus, the food-rich but colder and corrosive estuary induced a traits trade-off expressed as depressed IR under in situ conditions. However, this experience allowed these copepods to tolerate further exposure to high PCO2 levels better, as their IRs were on average 43% higher than those of the coastal individuals. Indeed, expression of both the Hsp70 and Ferritin genes in coastal copepods was significantly higher after acclimation to high PCO2 conditions. Along with other recent evidence, our findings confirm that adaptation to local fluctuations in seawater pH seems to play a significant role in the response of planktonic populations to OA-associated conditions. Facing the environmental threat represented by the inter-play between multiple drivers of climate change, this biological feature should be examined in detail as a potential tool for risk mitigation policies in coastal management arrangements.

Experiment: Ultraviolet radiation modulates the physiological responses of the calcified rhodophyte Corallina officinalis to elevated CO2

Ocean acidification reduces the concentration of carbonate ions and increases those of bicarbonate ions in seawater compared with the present oceanic conditions. This altered composition of inorganic carbon species may, by interacting with ultraviolet radiation (UVR), affect the physiology of macroalgal species. However, very little is known about how calcareous algae respond to UVR and ocean acidification. Therefore, we conducted an experiment to determine the effects of UVR and ocean acidification on the calcified rhodophyte Corallina officinalis using CO2-enriched cultures with and without UVR exposure. Low pH increased the relative electron transport rates (rETR) but decreased the CaCO3 content and had a miniscule effect on growth. However, UVA (4.25 W m-2) and a moderate level of UVB (0.5 W m-2) increased the rETR and growth rates in C. officinalis, and there was a significant interactive effect of pH and UVR on UVR-absorbing compound concentrations. Thus, at low irradiance, pH and UVR interact in a way that affects the multiple physiological responses of C. officinalis differently. In particular, changes in the skeletal content induced by low pH may affect how C. officinalis absorbs and uses light. Therefore, the light quality used in ocean acidification experiments will affect the predictions of how calcified macroalgae will respond to elevated CO2.