Uranium in larval shells as a barometer of molluscan ocean acidification exposure

As the ocean undergoes acidification, marine organisms will become increasingly exposed to reduced pH, yet variability in many coastal settings complicates our ability to accurately estimate pH exposure for those organisms that are difficult to track. Here we present shell-based geochemical proxies that reflect pH exposure from laboratory and field settings in larvae of the mussels Mytilus californianus and M. galloprovincialis. Laboratory-based proxies were generated from shells precipitated at pH 7.51 to 8.04. U/Ca, Sr/Ca, and multielemental signatures represented as principal components varied with pH for both species. Of these, U/Ca was the best predictor of pH and did not vary with larval size, with semidiurnal pH fluctuations, or with oxygen concentration. Field applications of U/Ca were tested with mussel larvae reared in situ at both known and unknown pH conditions. Larval shells precipitated in a region of greater upwelling had higher U/Ca, and these U/Ca values corresponded well with the laboratory-derived U/Ca-pH proxy. Retention of the larval shell after settlement in molluscs allows use of this geochemical proxy to assess ocean acidification effects on marine populations.

Combined effects of CO2 and temperature on carbon uptake and partitioning by the marine diatoms Thalassiosira weissflogii and Dactyliosolen fragilissimus

Carbon uptake and partitioning of two globally abundant diatom species, Thalassiosira weissflogii and Dactyliosolen fragilissimus, was investigated in batch culture experiments under four conditions: ambient (15°C, 400 µatm), high CO2 (15°C, 1000 µatm), high temperature (20°C, 400 µatm), and combined (20°C, 1000 µatm). The experiments were run from exponential growth into the stationary phase (six days after nitrogen depletion), allowing us to track biogeochemical dynamics analogous to bloom situations in the ocean. Elevated CO2 had a fertilizing effect and enhanced uptake of dissolved inorganic carbon (DIC) by about 8% for T. weissflogii and by up to 39% for D. fragilissimus. This was also reflected in higher cell numbers, build-up of particulate and dissolved organic matter, and transparent exopolymer particles. The CO2 effects were most prominent in the stationary phase when nitrogen was depleted and CO2(aq) concentrations were low. This indicates that diatoms in the high CO2 treatments could take up more DIC until CO2 concentrations in seawater became so low that carbon limitation occurs. These results suggest that, contrary to common assumptions, diatoms could be highly sensitive to ongoing changes in oceanic carbonate chemistry, particularly under nutrient limitation. Warming from 15 to 20 °C had a stimulating effect on one species but acted as a stressor on the other species, highlighting the importance of species-specific physiological optima and temperature ranges in the response to ocean warming. Overall, these sensitivities to CO2 and temperature could have profound impacts on diatoms blooms and the biological pump.

Colimitation of the unicellular photosynthetic diazotroph Crocosphaera watsonii by phosphorus, light, and carbon dioxide

We describe interactive effects of total phosphorus (total P = 0.1-4.0 µmol/L; added as H2NaPO4), irradiance (40 and 150 µmol quanta/m**2/s), and the partial pressure of carbon dioxide (P-CO2; 19 and 81 Pa, i.e., 190 and 800 ppm) on growth and CO2- and dinitrogen (N-2)-fixation rates of the unicellular N-2-fixing cyanobacterium Crocosphaera watsonii (WH0003) isolated from the Pacific Ocean near Hawaii. In semicontinuous cultures of C. watsonii, elevated P-CO2 positively affected growth and CO2- and N-2-fixation rates under high light. Under low light, elevated P-CO2 positively affected growth rates at all concentrations of P, but CO2- and N-2-fixation rates were affected by elevated P-CO2 only when P was low. In both high-light and low-light cultures, the total P requirements for growth and CO2- and N-2-fixation declined as P-CO2 increased. The minimum concentration (C-min) of total P and half-saturation constant (K-1/2) for growth and CO2- and N-2-fixation rates with respect to total P were reduced by 0.05 µmol/L as a function of elevated P-CO2. We speculate that low P requirements under high P-CO2 resulted from a lower energy demand associated with carbon-concentrating mechanisms in comparison with low-P-CO2 cultures. There was also a 0.10 µmol/L increase in C-min and K-1/2 for growth and N-2 fixation with respect to total P as a function of increasing light regardless of P-CO2 concentration. We speculate that cellular P concentrations are responsible for this shift through biodilution of cellular P and possibly cellular P uptake systems as a function of increasing light. Changing concentrations of P, CO2, and light have both positive and negative interactive effects on growth and CO2-, and N-2-fixation rates of unicellular oxygenic diazotrophs like C. watsonii.

CO2 and vitamin B12 interactions determine bioactive trace metal requirements of a subarctic Pacific diatom

Phytoplankton growth can be limited by numerous inorganic nutrients and organic growth factors. Using the subarctic diatom Attheya sp. in culture studies, we examined how the availability of vitamin B(12) and carbon dioxide partial pressure (pCO(2)) influences growth rate, primary productivity, cellular iron (Fe), cobalt (Co), zinc (Zn) and cadmium (Cd) quotas, and the net use efficiencies (NUEs) of these bioactive trace metals (mol C fixed per mol cellular trace metal per day). Under B(12)-replete conditions, cells grown at high pCO(2) had lower Fe, Zn and Cd quotas, and used those trace metals more efficiently in comparison with cells grown at low pCO(2). At high pCO(2), B(12)-limited cells had ~50% lower specific growth and carbon fixation rates, and used Fe ~15-fold less efficiently, and Zn and Cd ~3-fold less efficiently, in comparison with B(12)-replete cells. The observed higher Fe, Zn and Cd NUE under high pCO(2)/B(12)-replete conditions are consistent with predicted downregulation of carbon-concentrating mechanisms. Co quotas of B(12)-replete cells were 5- to 14-fold higher in comparison with B(12)-limited cells, suggesting that >80% of cellular Co of B(12)-limited cells was likely from B(12). Our results demonstrate that CO(2) and vitamin B(12) interactively influence growth, carbon fixation, trace metal requirements and trace metal NUE of this diatom. This suggests the need to consider complex feedback interactions between multiple environmental factors for this biogeochemically critical group of phytoplankton in the last glacial maximum as well as the current and future changing ocean.

Short- and long-term conditioning of a temperate marine diatom community to acidification and warming

Ocean acidification and greenhouse warming will interactively influence competitive success of key phytoplankton groups such as diatoms, but how long-term responses to global change will affect community structure is unknown. We incubated a mixed natural diatom community from coastal New Zealand waters in a short-term (two-week) incubation experiment using a factorial matrix of warming and/or elevated pCO2 and measured effects on community structure. We then isolated the dominant diatoms in clonal cultures and conditioned them for 1 year under the same temperature and pCO2 conditions from which they were isolated, in order to allow for extended selection or acclimation by these abiotic environmental change factors in the absence of interspecific interactions. These conditioned isolates were then recombined into 'artificial' communities modelled after the original natural assemblage and allowed to compete under conditions identical to those in the short-term natural community experiment. In general, the resulting structure of both the unconditioned natural community and conditioned 'artificial' community experiments was similar, despite differences such as the loss of two species in the latter. pCO2 and temperature had both individual and interactive effects on community structure, but temperature was more influential, as warming significantly reduced species richness. In this case, our short-term manipulative experiment with a mixed natural assemblage spanning weeks served as a reasonable proxy to predict the effects of global change forcing on diatom community structure after the component species were conditioned in isolation over an extended timescale. Future studies will be required to assess whether or not this is also the case for other types of algal communities from other marine regimes.

Chemical and physical oceanography on 131 CTD casts from METEOR cruise M59/2

The oceans absorb and store a significant portion of anthropogenic CO2 emissions, but large uncertainties remain in the quantification of this sink. An improved assessment of the present and future oceanic carbon sink is therefore necessary to provide recommendations for long-term global carbon cycle and climate policies. The formation of North Atlantic Deep Water (NADW) is a unique fast track for transporting anthropogenic CO2 into the ocean's interior, making the deep waters rich in anthropogenic carbon. Thus the Atlantic is presently estimated to hold 38% of the oceanic anthropogenic CO2 inventory, although its volume makes up only 25% of the world ocean. Here we analyze the inventory change of anthropogenic CO2 in the Atlantic between 1997 and 2003 and its relationship to NADW formation. For the whole region between 20°S and 65°N the inventory amounts to 32.5 ± 9.5 Petagram carbon (Pg C) in 1997 and increases up to 36.0 ± 10.5 Pg C in 2003. This result is quite similar to earlier studies. Moreover, the overall increase of anthropogenic carbon is in close agreement with the expected change due to rising atmospheric CO2 levels of 1.69% a?1. On the other hand, when considering the subpolar region only, the results demonstrate that the recent weakening in the formation of Labrador Sea Water, a component of NADW, has already led to a decrease of the anthropogenic carbon inventory in this water mass. As a consequence, the overall inventory for the total water column in the western subpolar North Atlantic increased only by 2% between 1997 and 2003, much less than the 11% that would be expected from the increase in atmospheric CO2 levels.

Physical oceanography from the TRACTOR project

Primary Objectives - Describe and quantify the present strength and variability of the circulation and oceanic processes of the Nordic Seas regions using primarily observations of the long term spread of a tracer purposefully released into the Greenland Sea Gyre in 1996. - Improve our understanding of ocean processes critical to the thermaholine circulation in the Nordic Seas regions so as to be able to predict how this region may respond to climate change. - Assess the role of mixing and ageing of water masses on the carbon transport and the role of the thermohaline circulation in carbon storage using water transports and mixing coefficients derived from the tracer distribution. Specific Objectives Perform annual hydrographic, chemical and SF6 tracer surveys into the Nordic regions in order to: - Measure lateral and diapycnal mixing rates in the Greenland Sea Gyre and in the surrounding regions. - Document the depth and rates of convective mixing in the Greenland Sea using the SF6 and the water masses characteristics. - Measure the transit time and transport of water from the Greenland Sea to surrounding seas and outflows. Document processes of water mass transformation and entrainment occurring to water emanating from the central Greenland Sea. - Measure diapycnal mixing rates in the bottom and margins of the Greenland Sea basin using the SF6 signal observed there. Quantify the potential role of bottom boundary-layer mixing in the ventilation of the Greenland Sea Deep Water in absence of deep convection. Monitor the variability of the entrainment of water from the Greenland Sea using time series auto-sampler moorings at strategic positions i.e., sill of the Denmark Strait, Labrador Sea, Jan Mayen fracture zone and Fram Strait. Relate the observed variability of the tracer signal in the outflows to convection events in the Greenland Sea and local wind stress events. Obtain a better description of deepwater overflow and entrainment processes in the Denmark Strait and Faeroe Bank Channel overflows and use these to improve modelling of deepwater overflows. Monitor the tracer invasion into the North Atlantic using opportunistic SF6 measurements from other cruises: we anticipate that a number of oceanographic cruises will take place in the north-east Atlantic and the Labrador Sea. It should be possible to get samples from some cruises for SF6 measurements. Use process models to describe the spread of the tracer to achieve better parameterisation for three-dimensional models. One reason that these are so resistant to prediction is that our best ocean models are as yet some distance from being good enough, to predict climate and climate change.