Seawater carbonate chemistry, primary production, biomass and calcification of plankton and bacteria, 2009

Production (abundance and biomass) and net calcification rates of the coccolithophorid Pleurochrysis carterae under different partial pressures of CO2 (pCO2) were examined using short (15, 24 and 39 h), long (7 d) and dark (7 d) incubation experiments. Short incubations were conducted at ambient, 500 and 820 ppm pCO2 levels in natural seawater that was enriched with nutrients and inoculated with P. carterae. Long incubations were conducted at ambient and 1200 ppm pCO2 levels in natural seawater (0.2 µm filtered as well as unfiltered) that was enriched with nutrients and inoculated with P. carterae. Dark incubations were conducted at ambient and 1200 ppm pCO2 in unfiltered seawater that was inoculated with P. carterae. The abundance and biomass of coccolithophorids increased with pCO2 and time. The abundance and biomass of most noncalcifying phytoplankton also increased, and were hardly affected by CO2 inputs. Net calcification rates were negative in short incubations during the pre-bloom phase regardless of pCO2 levels, indicating dissolution of calcium carbonate. Further, the negative values of net calcification in short incubations became less negative with time. Net calcification rates were positive in long incubations during blooms regardless of pCO2 level, and the rate of calcification increased with pCO2. Our results show that P. carterae may adapt to increased (~1200 ppm) pCO2 level with time, and such increase has little effect on the ecology of noncalcifying groups and hence in ecosystem dynamics. In dark incubations, net calcification rates were negative, with the magnitude being dependent on pCO2 levels.

Water chemistry, and abundance and carbon biomass of under-ice fauna during POLARSTERN cruise ARK-XIX/1 to the Storfjord area in 2003

The under-ice habitat and fauna were studied during a typical winter situation at three stations in the western Barents Sea. Dense pack ice (7-10/10) prevailed and ice thickness ranged over <0.1-1.6 m covered by <0.1-0.6 m of snow. Air temperatures ranged between -1.8 and -27.5°C. The ice undersides were level, white and smooth. Temperature and salinity profiles in the under-ice water (0-5 m depth) were not stratified (T=-1.9 to -2.0°C and S=34.2-34.7). Concentrations of inorganic nutrients were high and concentrations of algal pigments were very low (0.02 µg chlorophyll a/l), indicating the state of biological winter. Contents of particulate organic carbon and nitrogen ranged over 84.2-241.3 and 5.3-16.4 µg/l, respectively, the C/N ratio over 11.2-15.5 pointing to the dominance of detritus in the under-ice water. Abundances of amphipods at the ice underside were lower than in other seasons: 0-1.8 ind/m**2 for Apherusa glacialis, 0-0.7 ind/m**2 for Onisimus spp., and 0-0.8 ind/m**2 for Gammarus wilkitzkii. A total of 22 metazoan taxa were found in the under-ice water, with copepods as the most diverse and numerous group. Total abundances ranged over 181-2,487 ind/m**3 (biomass: 70-2,439 µg C/m**3), showing lower values than in spring, summer and autumn. The dominant species was the calanoid copepod Pseudocalanus minutus (34-1,485 ind/m**3), contributing 19-65% to total abundances, followed by copepod nauplii (85-548 ind/m**3) and the cyclopoid copepod Oithona similis (44-262 ind/m**3). Sympagic (ice-associated) organisms occurred only rarely in the under-ice water layer.

Seawater carbonate chemistry, biomass and calcification of Porites spp. corals during experiments, 2011

I tested the hypothesis that the effects of high pCO2 and temperature on massive Porites spp. (Scleractinia) are modified by heterotrophic feeding (zooplanktivory). Small colonies of massive Porites spp. from the back reef of Moorea, French Polynesia, were incubated for 1 month under combinations of temperature (29.3°C vs. 25.6°C), pCO2 (41.6 vs. 81.5 Pa), and feeding regimes (none vs. ad libitum access to live Artemia spp.), with the response assessed using calcification and biomass. Area-normalized calcification was unaffected by pCO2, temperature, and the interaction between the two, although it increased 40% with feeding. Biomass increased 35% with feeding and tended to be higher at 25.6°C compared to 29.3°C, and as a result, biomass-normalized calcification statistically was unaffected by feeding, but was depressed 12-17% by high pCO2, with the effect accentuated at 25.6°C. These results show that massive Porites spp. has the capacity to resist the effects on calcification of 1 month exposure to 81.5 Pa pCO2 through heterotrophy and changes in biomass. Area-normalized calcification is sustained at high pCO2 by a greater biomass with a reduced biomass-normalized rate of calcification. This mechanism may play a role in determining the extent to which corals can resist the long-term effects of ocean acidification.

Seawater carbonate chemistry, biomass and metabolic rates (leucine incorporation, CO2 fixation and respiration) of Rhodobacteraceae (strain MED165) and Flavobacteriaceae (strain MED217) in a laboratory experiment

Experimental results related to the effects of ocean acidification on planktonic marine microbes are still rather inconsistent and occasionally contradictory. Moreover, laboratory or field experiments that address the effects of changes in CO2 concentrations on heterotrophic microbes are very scarce, despite the major role of these organisms in the marine carbon cycle. We tested the direct effect of an elevated CO2 concentration (1000 ppmv) on the biomass and metabolic rates (leucine incorporation, CO2 fixation and respiration) of 2 isolates belonging to 2 relevant marine bacterial families, Rhodobacteraceae (strain MED165) and Flavobacteriaceae (strain MED217). Our results demonstrate that, contrary to some expectations, high pCO2 did not negatively affect bacterial growth but increased growth efficiency in the case of MED217. The elevated partial pressure of CO2 (pCO2) caused, in both cases, higher rates of CO2 fixation in the dissolved fraction and, in the case of MED217, lower respiration rates. Both responses would tend to increase the pH of seawater acting as a negative feedback between elevated atmospheric CO2 concentrations and ocean acidification.

Water chemistry, carbonate system parameters, and macro-organism biomass of vent and off-vent sites in the Columbretes Islands Marine Reserve

Ocean acidification is receiving increasing attention because of its potential to affect marine ecosystems. Rare CO2 vents offer a unique opportunity to investigate the response of benthic ecosystems to acidification. However, the benthic habitats investigated so far are mainly found at very shallow water (less than or equal to 5 m depth) and therefore are not representative of the broad range of continental shelf habitats. Here, we show that a decrease from pH 8.1 to 7.9 observed in a CO2 vent system at 40 m depth leads to a dramatic shift in highly diverse and structurally complex habitats. Forests of the kelp Laminaria rodriguezii usually found at larger depths (greater than 65 m) replace the otherwise dominant habitats (i.e. coralligenous outcrops and rhodolith beds), which are mainly characterized by calcifying organisms. Only the aragonite-calcifying algae are able to survive in acidified waters, while high-magnesium-calcite organisms are almost completely absent. Although a long-term survey of the venting area would be necessary to fully understand the effects of the variability of pH and other carbonate parameters over the structure and functioning of the investigated mesophotic habitats, our results suggest that in addition of significant changes at species level, moderate ocean acidification may entail major shifts in the distribution and dominance of key benthic ecosystems at regional scale, which could have broad ecological and socio-economic implications.

Water chemistry of lakes in Zackenbergdalen between 1997-2003, East Greenland

The ecology of arctic lakes is strongly influenced by climate-generated variations in snow coverage and by the duration of the ice-free period, which, in turn, affect the physical and chemical conditions of the lakes (Wrona et al., 2005, http://www.acia.uaf.edu/PDFs/ACIA_Science_Chapters_Final/ACIA_Ch08_Final.pdf). Most arctic lakes are characterised by a long period (8-10 months) of ice-cover, cold water and low algal biomass. The water temperature and nutrient concentrations, and most probably the nutrient input from the catchments, are closely related to the duration of snow- and ice-cover in the lakes. In years when the ice-out is late, - that is, in late July, - phytoplankton photosynthesis is limited by the lack of light and nutrients. Less food is then available to the next link in the food chain, such as copepods and daphnids, with implication on their growth rates.

Seawater carbonate chemistry in Ischia, Italy, 2008

The atmospheric partial pressure of carbon dioxide (pCO2) will almost certainly be double that of pre-industrial levels by 2100 and will be considerably higher than at any time during the past few million years1. The oceans are a principal sink for anthropogenic CO2 where it is estimated to have caused a 30% increase in the concentration of H+ in ocean surface waters since the early 1900s and may lead to a drop in seawater pH of up to 0.5 units by 2100. Our understanding of how increased ocean acidity may affect marine ecosystems is at present very limited as almost all studies have been in vitro, short-term, rapid perturbation experiments on isolated elements of the ecosystem4, 5. Here we show the effects of acidification on benthic ecosystems at shallow coastal sites where volcanic CO2 vents lower the pH of the water column. Along gradients of normal pH (8.1-8.2) to lowered pH (mean 7.8-7.9, minimum 7.4-7.5), typical rocky shore communities with abundant calcareous organisms shifted to communities lacking scleractinian corals with significant reductions in sea urchin and coralline algal abundance. To our knowledge, this is the first ecosystem-scale validation of predictions that these important groups of organisms are susceptible to elevated amounts of pCO2. Sea-grass production was highest in an area at mean pH 7.6 (1,827 µatm pCO2) where coralline algal biomass was significantly reduced and gastropod shells were dissolving due to periods of carbonate sub-saturation. The species populating the vent sites comprise a suite of organisms that are resilient to naturally high concentrations of pCO2 and indicate that ocean acidification may benefit highly invasive non-native algal species. Our results provide the first in situ insights into how shallow water marine communities might change when susceptible organisms are removed owing to ocean acidification.

Water chemistry and iron speciation in samples taken during SWEDARP 97/98 campaign with S.A. Agulhas to the Southern Ocean

The distribution and speciation of iron was determined along a transect in the eastern Atlantic sector (6°E) of the Southern Ocean during a collaborative Scandinavian/South African Antarctic cruise conducted in late austral summer (December 1997/January 1998). Elevated concentrations of dissolved iron (>0.4 nM) were found at 60°S in the vicinity of the Spring Ice Edge (SIE) in tandem with a phytoplankton bloom, chiefly dominated by Phaeocystis sp. This bloom had developed rapidly after the loss of the seasonal sea ice cover. The iron that fuelled this bloom was mostly likely derived from sea ice melt. In the Winter Ice Edge (WIE), around 55°S, dissolved iron concentrations were low (<0.2 nM) and corresponded to lower biological productivity, biomass. In the Antarctic Polar Front, at approximately 50°S, a vertical profile of dissolved iron showed low concentrations (<0.2 nM); however, a surface survey showed higher concentrations (1-3 nM), and considerable patchiness in this dynamic frontal region. The chemical speciation of iron was dominated by organic complexation throughout the study region. Organic iron-complexing ligands ([L]) ranged from 0.9 to 3.0 nM Fe equivalents, with complex stability log K'(FeL) = 21.4-23.5. Estimated concentrations of inorganic iron (Fe') ranged from 0.03 to 0.79 pM, with the highest values found in the Phaeocystis bloom in the SIE. A vertical profile of iron-complexing ligands in the WIE showed a maximum consistent with a biological source for ligand production and near surface minimum possibly consistent with loss via photodecomposition. This work further confirms the role iron that has in the Southern Ocean in limiting primary productivity.

Ladderane lipid analysis and pore water and sediment chemistry of sediment cores from the continental shelf and slope off northwest Africa

The presence and abundance of anaerobic ammonium-oxidizing (anammox) bacteria was investigated in continental shelf and slope sediments (300-3000 m water depth) off northwest Africa in a combined approach applying quantitative polymerase chain reaction (q-PCR) analysis of anammox-specific 16S rRNA genes and anammox-specific ladderane biomarker lipids. We used the presence of an intact ladderane monoether lipid with a phosphocholine (PC) headgroup as a direct indicator for living anammox bacteria and compared it with the abundance of ladderane core lipids derived from both living and dead bacterial biomass. All investigated sediments contained ladderane lipids, both intact and core lipids, in agreement with the presence of anammoxspecific 16S rRNA gene copies, indicating that anammox occurs at all sites. Concentrations of ladderane core lipids in core top sediments varied between 0.3 and 97 ng g**-1 sediment, with the highest concentrations detected at the sites located on the shelf at shallower water depths between 300 and 500 m. In contrast, the C20 [3]-ladderane monoether-PC lipid was most abundant in a core top sediment from 1500 m water depth. Both anammox-specific 16S rRNA gene copy numbers and the concentration of the C20 [3]-ladderane monoether-PC lipid increased downcore in sediments located at greater water depths, showing highest concentrations of 1.2 x 10**8 copies g**-1 sediment and 30 pg g**-1 sediment, respectively, at the deepest station of 3000 m water depth. This suggests that the relative abundance of anammox bacteria is higher in sediments at intermediate to deep water depths where carbon mineralization rates are lower but where anammox is probably more important than denitrification.

Seawater carbonate chemistry, pigments and proteins during experiments with phytoplankton, 2010

In the high-nutrient, low-chlorophyll waters of the Gulf of Alaska, microcosm manipulation experiments were used to assess the effect of CO2 on growth and primary production under iron-limited and iron-replete conditions. As expected, iron had a strong effect on growth and photosynthesis. A modest and variable stimulation of growth and biomass production by CO2 (high CO2: 77-122 Pa; low CO2: 11-17 Pa) was observed under both iron-replete and iron-limited conditions, though near the limit of precision of our measurements in slow-growing low-iron experiments. Physiological acclimations responsible for the changes in growth were assessed. Under iron-limited conditions, growth stimulation at high CO2 appeared to result from an increase in photosynthetic efficiency, which we attribute to energy savings from down-regulation of the carbon concentrating mechanisms. In some cases, iron-rich photosynthetic proteins (PsbA, PsaC, and cytochrome b6) were down-regulated at elevated CO2in iron-limited controls. Under iron-replete conditions, there was an increase in growth rate and biomass at high CO2 in some experiments. This increase was unexpectedly supported by reductions in cellular carbon loss, most likely decreased respiration. We speculate that this effect may be due to acclimation to decreased pH rather than high CO2. The variability in responses to CO2 among experiments did not appear to be caused by differences in phytoplankton community structure and may reflect the sensitivity of the net response of phytoplankton to antagonistic effects of the several parameters that co-vary with CO2.

Seawater carbonate chemistry and processes during experiments with cyanobacterium Nodularia spumigena, 2009

The surface ocean absorbs large quantities of the CO2 emitted to the atmosphere from human activities. As this CO2 dissolves in seawater, it reacts to form carbonic acid. While this phenomenon, called ocean acidification, has been found to adversely affect many calcifying organisms, some photosynthetic organisms appear to benefit from increasing [CO2]. Among these is the cyanobacterium Trichodesmium, a predominant diazotroph (nitrogen-fixing) in large parts of the oligotrophic oceans, which responded with increased carbon and nitrogen fixation at elevated pCO2. With the mechanism underlying this CO2 stimulation still unknown, the question arises whether this is a common response of diazotrophic cyanobacteria. In this study we therefore investigate the physiological response of Nodularia spumigena, a heterocystous bloom-forming diazotroph of the Baltic Sea, to CO2-induced changes in seawater carbonate chemistry. N. spumigena reacted to seawater acidification/carbonation with reduced cell division rates and nitrogen fixation rates, accompanied by significant changes in carbon and phosphorus quota and elemental composition of the formed biomass. Possible explanations for the contrasting physiological responses of Nodularia compared to Trichodesmium may be found in the different ecological strategies of non-heterocystous (Trichodesmium) and heterocystous (Nodularia) cyanobacteria.

Seawater carbonate chemistry and calcification during tropical reef studies at Moorea (French Polynesia), 1997

Community metabolism and air-sea carbon dioxide (CO2) fluxes were investigated in July 1992 on a fringing reef at Moorea (French Polynesia). The benthic community was dominated by macroalgae (85% substratum cover) and comprised of Phaeophyceae Padina tenuis (Bory), Turbinaria ornata (Turner) J. Agardh, and Hydroclathrus clathratus Bory (Howe); Chlorophyta Halimeda incrassata f. ovata J. Agardh (Howe); and Ventricaria ventricosa J. Agardh (Olsen et West), as well as several Rhodophyta (Actinotrichia fragilis Forskál (Børgesen) and several species of encrusting coralline algae). Algal biomass was 171 g dry weight/m**2. Community gross production (Pg), respiration (R), and net calcification (G) were measured in an open-top enclosure. Pg and R were respectively 248 and 240 mmol Co2/m**2/d, and there was a slight net dissolution of CaCO3 (0.8 mmol/m**2/d). This site was a sink for atmospheric CO2 (10 ± 4 mmol CO2/m**2/d), and the analysis of data from the literature suggests that this is a general feature of algal-dominated reefs. Measurement of air-sea CO2 fluxes in open water close to the enclosure demonstrated that changes in small-scale hydrodynamics can lead to misleading conclusions. Net CO2 evasion to the atmosphere was measured on the fringing reef due to changes in the current pattern that drove water from the barrier reef (a C02 source) to the study site.

(Table 1) Near surface water chemistry of south Patagonian waters during spring and winter

Both the biomass of autotrophic dinoflagellates and its contribution to total chlorophyll were found to increase significantly with seawater temperature and the level of stratification in southern Patagonian waters during spring and winter. The highest peak of biomass corresponded to a single species, Prorocentrum minimum (Pavillard) Schiller, and was detected in middle shelf waters, coinciding with the primary productivity and CO2 uptake maxima reported for the area under spring conditions.