Tidal downwelling and implications for the carbon biogeochemistry of cold-water corals in relation to future ocean acidification and warming

Cold-water coral (CWC) reefs are recognised as ecologically and biologically significant areas that generate habitats and diversity. The interaction between hydrodynamics and CWCs has been well-studied at the Mingulay Reef Complex, a relatively shallow area of reefs found on the continental shelf off Scotland, UK. Within ‘Mingulay Area 01’ a rapid tidal downwelling of surface waters, brought about as an internal wave, is known to supply warmer, phytoplankton-rich waters to corals growing on the northern flank of an east-west trending seabed ridge. This study shows that this tidal downwelling also causes short-term perturbations in the inorganic carbon and nutrient dynamics through the water column and immediately above the reef. Over a 14 h period, corresponding to one semi-diurnal tidal cycle, seawater pH overlying the reef varied by ~0.1 pH unit, while pCO₂ shifted by > 60 μatm, a shift equivalent to a ~25 year jump into the future, with respect to atmospheric pCO₂. During the summer stratified period, these downwelling events result in the reef being washed over with surface water that has higher pH, is warmer, nutrient-depleted, but rich in phytoplankton-derived particles compared to the deeper waters in which the corals sit. Empirical observations, together with outputs from the European Regional Shelf Sea Ecosystem Model, demonstrate that the variability that the CWC reefs experience changes through the seasons and into the future. Hence, as ocean acidification and warming increase into the future, the downwelling event specific to this site could provide short-term amelioration of corrosive conditions at certain times of the year; however it could additionally result in enhanced detrimental impacts of warming on CWCs. Natural variability in the inorganic carbon and nutrient conditions, as well as local hydrodynamic regimes, must be accounted for in any future predictions concerning the responses of marine ecosystems to climate change.

The effect of ocean acidification on early algal colonization stages at natural CO2 vents

Marine algae exhibit different responses to ocean acidification, suggesting that a decrease in pH does not always favour marine photosynthetic organisms. In order to understand the effect of acidification on algal community development, early colonization stages were investigated using carbon dioxide vents around the Castello Aragonese (Ischia, Italy) as a natural laboratory. Settlement tiles were placed in zones with different pH (normal, medium and low), and species composition and coverage measured after 2, 3 and 4 months of deployment. The number of species decreased by 4 and 18 % at medium and low pH zones, respectively (P < 0.05). The structure of the algal assemblage differed between pH zones during the 4 months of the experiment, due to the addition and/or replacement of new species. This leads to a change in the succession of morphological forms as soft crustose algae replaced calcareous species, and turf species were dominant in cover; more complex thalli started to occur only at medium pH. These results support previous findings that ocean acidification will induce changes in benthic algal communities.

Arctic rhodolith beds and their environmental controls (Spitsbergen, Norway)

Coralline algae (Corallinales, Rhodophyta) that form rhodoliths are important ecosystem engineers and carbonate producers in many polar coastal habitats. This study deals with rhodolith communities from Floskjeret (78°18′N), Krossfjorden (79°08′N), and Mosselbukta (79°53′N), off Spitsbergen Island, Svalbard Archipelago, Norway. Strong seasonal variations in temperature, salinity, light regime, sea-ice coverage, and turbidity characterize these localities. The coralline algal flora consists of Lithothamnion glaciale and Phymatolithon tenue. Well-developed rhodoliths were recorded between 27 and 47 m water depth, while coralline algal encrustations on lithoclastic cobbles were detected down to 77 m water depth. At all sites, ambient waters were saturated with respect to both aragonite and calcite, and the rhodolith beds were located predominately at dysphotic water depths. The rhodolith-associated macrobenthic fauna included grazing organisms such as chitons and echinoids. With decreasing water depth, the rhodolith pavements were regularly overgrown by non-calcareous Polysiphonia-like red algae. The corallines are thriving and are highly specialized in their adaptations to the physical environment as well as in their interaction with the associated benthic fauna, which is similar to other polar rhodolith communities. The marine environment of Spitsbergen is already affected by a climate-driven ecological regime shift and will lead to an increased borealization in the near future, with presently unpredictable consequences for coralline red algal communities.

Occurrence of a cold-water coral along natural pH gradients (Patagonia, Chile)

Increasing dissolution of anthropogenic-released carbon dioxide into the world’s oceans is causing ocean acidification (OA). OA is thought to negatively affect most marine-calcifying organisms, notably cold-water corals (CWC), which may be especially sensitive due to the deep and cold waters they normally thrive in. However, the impact of OA on CWC is difficult to predict. Recorded distributions of CWC are rarely linked to in situ water chemistry, and the boundaries of their distributions are not clearly defined. The fjord Comau in Chilean Patagonia features pronounced pH gradients, and up to 0.5 pH units have been recorded both vertically (at some sites within 50 m depth) and less distinct horizontally (from head to mouth). The cosmopolite coral Desmophyllum dianthus grows along the course of the fjord and of the entire pH range. It occurs in shallow depths (below 12 m, pH 8.1) as part of a deep-water emergence community, but also in 225 m depth at a pH of 7.4. Based on pH and total alkalinity, data calculations of the associated carbonate chemistry revealed that this CWC thrives commonly close the aragonite (the orthogonal crystal form of calcium carbonate, the mineral structure of coral skeletons) saturation horizon and even below. This suggests a high adaptation potential of D. dianthus to adjust its calcification performance to conditions thermodynamically unfavourable for the precipitation of aragonite.

Results of laboratory and field experiments of the direct effect of increasing CO2 on net primary production of macroalgal species in brackish-water ecosystems

Studies on the effects of increasing acidification on marine communities have been previously mostly carried out in truly marine areas whereas brackish-water ecosystems such as the Baltic Sea have been less studied. The current study analyses how acidification induced by elevated atmospheric carbon dioxide affects the photosynthetic net production of different macroalgal species in the brackish Baltic Sea. Research methods include sets of laboratory and field experiments carried out in shallow coastal brackish waters. The aim of the laboratory experiments was to develop the necessary techniques and experience for the mesocosm experiments. Laboratory experiments were carried out using specimens of the red alga Furcellaria lumbricalis collected from Kakumäe Bay. The mesocosm experiments were conducted in Kõiguste Bay during the field season of 2011. Separate mesocosms were operated in each set with different CO₂ concentrations and a control treatment in natural conditions. Field experiments were carried out with three species representing three different morphological and ecological groups: Ulva intestinalis, a fast-growing green alga; Fucus vesiculosus, a perennial brown alga with a slow metabolism; and Furcellaria lumbricalis, a perennial red alga. Photosynthetic activity was used as the response variable. In the laboratory decreasing pH increased the net primary production of F. lumbricalis with the lowest net primary production values measured at pH 8.0 and the highest at pH 6.5. Results of the field experiments indicated that increased CO₂ levels in seawater favoured photosynthetic activity of the macroalgae U. intestinalis and F. lumbricalis, but F. vesiculosus showed no response to elevated CO₂. Elevated CO₂ levels are suggested to favour the production of fast-growing filamentous species, which thus may indirectly enhance the effect of eutrophication in the shallow coastal brackish waters.

Development of a colorimetric microfluidic pH sensor for autonomous seawater measurements

High quality carbonate chemistry measurements are required in order to fully understand the dynamics of the oceanic carbonate system. Seawater pH data with good spatial and temporal coverage are particularly critical to apprehend ocean acidification phenomena and their consequences. There is a growing need for autonomous in situ instruments that measure pH on remote platforms. Our aim is to develop an accurate and precise autonomous in situ pH sensor for long term deployment on remote platforms. The widely used spectrophotometric pH technique is capable of the required high-quality measurements. We report a key step toward the miniaturisation of a colorimetric pH sensor with the successful implementation of a simple microfluidic design with low reagent consumption. The system is particularly adapted to shipboard deployment: high quality data was obtained over a period of more than a month during a shipboard deployment in northwest European shelf waters, and less than 30 mL of indicator was consumed. The system featured a short term precision of 0.001 pH (n=20) and an accuracy within the range of a certified Tris buffer (0.004 pH). The quality of the pH system measurements have been checked using various approaches: measurements of certified Tris buffer, measurement of certified seawater for DIC and TA, comparison of measured pH against calculated pH from pCO₂, DIC and TA during the cruise in northwest European shelf waters. All showed that our measurements were of high quality. The measurements were made close to in situ temperature (+0.2 °C) in a sampling chamber which had a continuous flow of the ship’s underway seawater supply. The optical set up was robust and relatively small due to the use of an USB mini-spectrometer, a custom made polymeric flow cell and an LED light source. The use of a three wavelength LED with detection that integrated power across the whole of each LED output spectrum indicated that low wavelength resolution detectors can be used instead of the current USB mini spectrophotometer. Artefacts due to the polychromatic light source and inhomogeneity in the absorption cell are shown to have a negligible impact on the data quality. The next step in the miniaturisation of the sensor will be the incorporation of a photodiode as detector to replace the spectrophotometer.

Assessing seasonal changes in carbonate parameters across small spatial gradients in the Northeastern Chukchi Sea

Observations of the marine carbonate system were made in 2010 in the northeastern Chukchi Sea to constrain the seasonal progression of carbonate mineral saturation states (Ω) throughout the water column and determine the air-sea flux of carbon dioxide (CO₂). As sea ice retreats from the Chukchi Shelf, primary production consumes dissolved inorganic carbon (DIC) in the euphotic zone causing pH and carbonate mineral saturation states to increase. Throughout the summer and early autumn months of 2010, saturation states for calcite and aragonite ranged from 2.5–4.0 and 1.5–2.5, respectively, well about the saturation horizon of 1.0. Much of the organic matter produced during the bloom was vertically exported from the relatively small study area leading to an uptake of CO₂ from the atmosphere of at least 340,000 kg-C. The exported organic matter settled near the bottom and was remineralized back into DIC, causing concentrations to increase sharply, particularly in autumn months, driving down pH to as low as 7.75 and suppressing the concentrations of important carbonate minerals to the point that aragonite became undersaturated. The data showed a definitive seasonal progression of this process with aragonite becoming partially undersaturated along the bottom in September, and broadly undersaturated in October. While carbonate saturation states would naturally be suppressed by the high rates of export production and the accumulation of DIC near the bottom, the penetration of anthropogenic CO₂ into water column (ocean acidification) has caused these observed undersaturations, which will likely expand as CO₂ levels in the atmosphere continue to rise in the coming decades.

Ocean acidification from 1997 to 2011 in the subarctic western North Pacific Ocean

Rising atmospheric CO₂ contents have led to greater CO₂ uptake by the oceans, lowering both pH due to increasing hydrogen ions and CaCO₃ saturation states due to declining carbonate ion (CO₃²⁻). Here, we used previously compiled data sets and new data collected in 2010 and 2011 to investigate ocean acidification of the North Pacific western subarctic gyre. In winter, the western subarctic gyre is a source of CO₂ to the atmosphere because of convective mixing of deep waters rich in dissolved inorganic carbon (DIC). We calculated pH in winter mixed layer from DIC and total alkalinity (TA), and found that it decreased at the rate of −0.001 ± 0.0004 yr⁻¹ from 1997 to 2011. This decrease rate is slower than that expected under condition of seawater/atmosphere equilibration, and it is also slower than the rate in the subtropical regions (−0.002 yr⁻¹). The slow rate is caused by a reduction of CO₂ emission in winter due to an increase in TA. Below the mixed layer, the calcite saturation horizon (~185 m depth) shoaled at the rate of 2.9 ± 0.9 m yr⁻¹ as the result of the declining CO₃²⁻ concentration (−0.03 ± 0.01 μmol k⁻¹yr⁻¹). Between 200 m and 300 m depth, pH decline during the study period (−0.0051 ± 0.0010 yr⁻¹) was larger than ever reported in the open North Pacific. This enhanced acidification rate below the calcite saturation horizon reflected not only the uptake of anthropogenic CO₂ but also the increase in the decomposition of organic matter evaluated from the increase in AOU, which suggests that the dissolution of CaCO₃ particles increased.

Effect of CO2 enrichment on bacterial metabolism in an Arctic fjord (update)

The anthropogenic increase of carbon dioxide (CO₂) alters the seawater carbonate chemistry, with a decline of pH and an increase in the partial pressure of CO₂ (pCO₂). Although bacteria play a major role in carbon cycling, little is known about the impact of rising pCO₂ on bacterial carbon metabolism, especially for natural bacterial communities. In this study, we investigated the effect of rising pCO₂ on bacterial production (BP), bacterial respiration (BR) and bacterial carbon metabolism during a mesocosm experiment performed in Kongsfjorden (Svalbard) in 2010. Nine mesocosms with pCO₂ levels ranging from ca. 180 to 1400 μatm were deployed in the fjord and monitored for 30 days. Generally BP gradually decreased in all mesocosms in an initial phase, showed a large (3.6-fold average) but temporary increase on day 10, and increased slightly after inorganic nutrient addition. Over the wide range of pCO₂ investigated, the patterns in BP and growth rate of bulk and free-living communities were generally similar over time. However, BP of the bulk community significantly decreased with increasing pCO₂ after nutrient addition (day 14). In addition, increasing pCO₂ enhanced the leucine to thymidine (Leu : TdR) ratio at the end of experiment, suggesting that pCO₂ may alter the growth balance of bacteria. Stepwise multiple regression analysis suggests that multiple factors, including pCO₂, explained the changes of BP, growth rate and Leu : TdR ratio at the end of the experiment. In contrast to BP, no clear trend and effect of changes of pCO₂ was observed for BR, bacterial carbon demand and bacterial growth efficiency. Overall, the results suggest that changes in pCO₂ potentially influence bacterial production, growth rate and growth balance rather than the conversion of dissolved organic matter into CO₂.

Ocean acidification state in western Antarctic surface waters: drivers and interannual variability

Each December during four years from 2006 to 2010, the surface water carbonate system was measured and investigated in the Amundsen Sea and Ross Sea, western Antarctica as part of the Oden Southern Ocean expeditions (OSO). The I/B Oden started in Punta Arenas in Chile and sailed southwest, passing through different regimes such as, the marginal/seasonal ice zone, fronts, coastal shelves, and polynyas. Discrete surface water was sampled underway for analysis of total alkalinity (A_T), total dissolved inorganic carbon (C_T) and pH. Two of these parameters were used together with sea-surface temperature (SST), and salinity to obtain a full description of the surface water carbonate system, including pH in situ and calcium carbonate saturation state of aragonite (Ω_Ar) and calcite (Ω_Ca). Multivariate analysis was used to investigate interannual variability and the major controls (sea-ice concentration, SST, salinity and chlorophyll a) on the variability in the carbonate system and Ω. This analysis showed that SST and chlorophyll a were the major drivers of the Ω variability in both the Amundsen and Ross seas. In 2007, the sea-ice edge was located further south and the area of the open polynya was relatively small compared to 2010. We found the lowest pH in situ (7.932) and Ω = 1 values in the sea-ice zone and in the coastal Amundsen Sea, nearby marine out flowing glaciers. In 2010, the sea-ice coverage was the largest and the areas of the open polynyas were the largest for the whole period. This year we found the lowest salinity and A_T, coinciding with highest chl a. This implies that the highest Ω_Ar in 2010 was likely an effect of biological CO₂ drawdown, which out-competed the dilution of carbonate ion concentration due to large melt water volumes. We predict and discuss future Ω values, using our data and reported rates of oceanic uptake of anthropogenic CO₂, suggesting that the Amundsen Sea will become undersaturated with regard to aragonite about 20 yr sooner than predicted by models.