Seawater carbonate chemistry and its effects on properties and sinking velocity of aggregates of the coccolithophore Emiliania huxleyi, 2010

Coccolithophores play an important role in organic matter export due to their production of the mineral calcite that can act as ballast. Recent studies indicated that calcification in coccolithophores may be affected by changes in seawater carbonate chemistry. We investigated the influence of CO2 on the aggregation and sinking behaviour of the coccolithophore Emiliania huxleyi (PML B92/11) during a laboratory experiment. The coccolithophores were grown under low (~180 µatm), medium (~380 µatm), and high (~750 µatm) CO2 conditions. Aggregation of the cells was promoted using roller tables. Size and settling velocity of aggregates were determined during the incubation using video image analysis. Our results indicate that aggregate properties are sensitive to changes in the degree of ballasting, as evoked by ocean acidification. Average sinking velocity was highest for low CO2 aggregates (~1292 m d-1) that also had the highest particulate inorganic to particulate organic carbon (PIC/POC) ratio. Lowest PIC/POC ratios and lowest sinking velocity (~366 m d-1) at comparable sizes were observed for aggregates of the high CO2 treatment. Aggregates of the high CO2 treatment showed a 4-fold lower excess density (~4.2*10**-4 g cm**-3) when compared to aggregates from the medium and low CO2 treatments (~1.7 g*10**-3 cm**-3). We also observed that more aggregates formed in the high CO2 treatment, and that those aggregates contained more bacteria than aggregates in the medium and low CO2 treatment. If applicable to the future ocean, our findings suggest that a CO2 induced reduction of the calcite content of aggregates could weaken the deep export of organic matter in the ocean, particularly in areas dominated by coccolithophores.

Seawater carbonate chemistry, photosynthesis, respiration and calcification during experiments with scleractinian coral Stylophora pistillata, 2003

We show here that CO2 partial pressure (pCO2) and temperature significantly interact on coral physiology. The effects of increased pCO2 and temperature on photosynthesis, respiration and calcification rates were investigated in the scleractinian coral Stylophora pistillata. Cuttings were exposed to temperatures of 25°C or 28°C and to pCO2 values of ca. 460 or 760 muatm for 5 weeks. The contents of chlorophyll c2 and protein remained constant throughout the experiment, while the chlorophyll a content was significantly affected by temperature, and was higher under the 'high-temperature-high-pCO2' condition. The cell-specific density was higher at 'high pCO2' than at 'normal pCO2' (1.7 vs. 1.4). The net photosynthesis normalized per unit protein was affected by both temperature and pCO2, whereas respiration was not affected by the treatments. Calcification decreased by 50% when temperature and pCO2 were both elevated. Calcification under normal temperature did not change in response to an increased pCO2. This is not in agreement with numerous published papers that describe a negative relationship between marine calcification and CO2. The confounding effect of temperature has the potential to explain a large portion of the variability of the relationship between calcification and pCO2 reported in the literature, and warrants a re-evaluation of the projected decrease of marine calcification by the year 2100.

Coral and mollusc resistance to ocean acidification adversely affected by warming, 2011

Increasing atmospheric carbon dioxide (CO2) concentrations are expectedto decrease surface ocean pH by 0.3-0.5 units by 2100, lowering the carbonate ion concentration of surfacewaters. This rapid acidification is predicted to dramatically decrease calcification in many marine organisms. Reduced skeletal growth under increased CO2 levels has already been shown for corals, molluscs and many other marine organisms. The impact of acidification on the ability of individual species to calcify has remained elusive, however, as measuring net calcification fails to disentangle the relative contributions of gross calcification and dissolution rates on growth. Here, we show that corals and molluscs transplanted along gradients of carbonate saturation state at Mediterranean CO2 vents are able to calcify and grow at even faster than normal rates when exposed to the high CO2 levels projected for the next 300 years. Calcifiers remain at risk, however, owing to the dissolution of exposed shells and skeletons that occurs as pH levels fall. Our results show that tissues and external organic layers play a major role in protecting shells and skeletons from corrosive sea water, limiting dissolution and allowing organisms to calcify. Our combined field and laboratory results demonstrate that the adverse effects of global warming are exacerbated when high temperatures coincide with acidification.

Seawater carbonate chemistry and echinoderm Asterias rubens biological processes during experiments, 2011

We compared effects of exposure to predicted near-future (2100) ocean acidification (OA; pH 7.7) and normal seawater (Control; pH 8.1) on immune and stress responses in the adult sea star Asterias rubens. Analyses were made after one week and after six months of continuous exposure. Following one week exposure to acidified water, the pH of coelomic fluid was significantly reduced. Levels of the chaperon Hsp70 were elevated while key cellular players in immunity, coelomocytes, were reduced by approximately 50%. Following long-term exposure (six months) levels of Hsp70 returned to control values, whereas immunity was further impaired, evidenced by the reduced phagocytic capacity of coelomocytes and inhibited activation of p38 MAP-kinase. Such impacts of reduced seawater pH may have serious consequences for resistance to pathogens in a future acidified ocean.

Seawater carbonate chemistry, chlorophyll a and phosphate during experiments with Trichodesmium erythraeum IMS101 (CCMP1985), 2010

In this laboratory study, we monitored the buildup of biomass and concomitant shift in seawater carbonate chemistry over the course of a Trichodesmium bloom under different phosphorus (P) availability. During exponential growth, dissolved inorganic carbon (DIC) decreased, while pH increased until maximum cell densities were reached. Once P became depleted, DIC decreased even further and total alkalinity (TA) dropped, accompanied by precipitation of aragonite. Under P-replete conditions, DIC increased and TA remained constant in the postbloom phase. A diffusion-reaction model was employed to estimate changes in carbonate chemistry of the diffusive boundary layer. This study demonstrates that Trichodesmium can induce precipitation of aragonite from seawater and further provides possible explanations about underlying mechanisms.

Seawater carbonate chemistry and crab Necora puber size and elements in tissue during experiments, 2010

Ocean acidification (OA) is predicted to play a major role in shaping species biogeography and marine biodiversity over the next century. We tested the effect of medium-term exposure to OA (pH 8.00, 7.30 and 6.70 for 30 d) on acid-base balance in the decapod crab Necora puber-a species that is known to possess good extracellular buffering ability during short-term exposure to hypercapnic conditions. To determine if crabs undergo physiological trade-offs in order to buffer their haemolymph, we characterised a number of fundamental physiological functions, i.e. metabolic rate, tolerance to heat, carapace and chelae [Ca2+] and [Mg2+], haemolymph [Ca2+] and [Mg2+], and immune response in terms of lipid peroxidation. Necora puber was able to buffer changes to extracellular pH over 30 d exposure to hypercapnic water, with no evidence of net shell dissolution, thus demonstrating that HCO3- is actively taken up from the surrounding water. In addition, tolerance to heat, carapace mineralization, and aspects of immune response were not affected by hypercapnic conditions. In contrast, whole-animal O2uptake significantly decreased with hypercapnia, while significant increases in haemolymph [Ca2+] and [Mg2+] and chelae [Mg2+] were observed with hypercapnia. Our results confirm that most physiological functions in N. puber are resistant to low pH/hypercapnia over a longer period than previously investigated, although such resistance comes at the expenses of metabolic rates, haemolymph chemistry and chelae mineralization.

Seawater carbonate chemistry and biological processes during experiments with Patella vulgata, 2010

The effect of short-term (5 days) exposure to CO2-acidified seawater (year 2100 predicted values, ocean pH = 7.6) on key aspects of the function of the intertidal common limpet Patella vulgata (Gastropoda: Patellidae) was investigated. Changes in extracellular acid-base balance were almost completely compensated by an increase in bicarbonate ions. A concomitant increase in haemolymph Ca2+ and visible shell dissolution implicated passive shell dissolution as the bicarbonate source. Analysis of the radula using SEM revealed that individuals from the hypercapnic treatment showed an increase in the number of damaged teeth and the extent to which such teeth were damaged compared with controls. As radula teeth are composed mainly of chitin, acid dissolution seems unlikely, and so the proximate cause of damage is unknown. There was no hypercapnia-related change in metabolism (O2 uptake) or feeding rate, also discounting the possibility that teeth damage was a result of a CO2-related increase in grazing. We conclude that although the limpet appears to have the physiological capacity to maintain its extracellular acid-base balance, metabolism and feeding rate over a 5 days exposure to acidified seawater, radular damage somehow incurred during this time could still compromise feeding in the longer term, in turn decreasing the top-down ecosystem control that P. vulgata exerts over rocky shore environments.

Seawater carbonate chemistry and biological processes of coccolithophore (Gephyrocapsa oceanica and Coccolithus pelagicus ssp. braarudii) during experiments, 2011

All species of coccolithophore appear to respond to perturbations of carbonate chemistry in a different way. Here, we show that the degree of malformation, growth rate and stable isotopic composition of organic matter and carbonate produced by two contrasting species of coccolithophore (Gephyrocapsa oceanica and Coccolithus pelagicus ssp. braarudii) are indicative of differences between their photosynthetic and calcification response to changing DIC levels (ranging from ~1100 to ~7800 µmol/kg) at constant pH (8.13 ± 0.02). Gephyrocapsa oceanica thrived under all conditions of DIC, showing evidence of increased growth rates at higher DIC, but C. braarudii was detrimentally affected at high DIC showing signs of malformation, and decreased growth rates. The carbon isotopic fractionation into organic matter and the coccoliths suggests that C. braarudii utilises a common internal pool of carbon for calcification and photosynthesis but G. oceanica relies on independent supplies for each process. All coccolithophores appear to utilize bicarbonate as their ultimate source of carbon for calcification resulting in the release of a proton. But, we suggest that this proton can be harnessed to enhance the supply of CO2(aq) for photosynthesis either from a large internal HCO3- pool which acts as a pH buffer (C. braarudii), or pumped externally to aid the diffusive supply of CO2 across the membrane from the abundant HCO3- (G. oceanica), likely mediated by an internal and external carbonic anhydrase respectively. Our simplified hypothetical spectrum of physiologies may provide a context to understand different species response to changing pH and DIC, the species-specific delta p and calcite "vital effects", as well as accounting for geological trends in coccolithophore cell size.

Seawater carbonate chemistry and calcification during experiments with corals, 2003

Biogenic calcification is influenced by the concentration of available carbonate ions. The recent confirmation of this for hermatypic corals has raised concern over the future of coral reefs because [CO3] is a decreasing function of increasing pCO2 in the atmosphere. As one of the overriding features of coral reefs is their diversity, understanding the degree of variability between species in their ability to cope with a change in [CO3] is a priority. We cultured four phylogenetically and physiologically different species of hermatypic coral (Acropora verweyi, Galaxea fascicularis, Pavona cactus and Turbinaria reniformis) under 'normal' (280 µmol/kg) and 'low' (140 µmol/kg) carbonate-ion concentrations. The effect on skeletogenesis was investigated quantitatively (by calcification rate) and qualitatively (by microstructural appearance of growing crystalline fibres using scanning electron microscopy (SEM)). The 'low carbonate' treatment resulted in a significant suppression of calcification rate and a tendency for weaker crystallization at the distal tips of fibres. However, while the calcification rate was affected uniformly across species (13-18% reduction), the magnitude of the microstructural response was highly species specific: crystallization was most markedly affected in A. verweyi and least in T. reniformis. These results are discussed in relation to past records and future predictions of carbonate variability in the oceans.

Effects of ocean acidification on the calcification of otoliths of larval Atlantic cod Gadus morhua

The growth and development of the aragonitic CaCO3 otoliths of teleost fish could be vulnerable to processes resulting from ocean acidification. The potential effects of an increase in atmospheric CO2 on the calcification of the otoliths were investigated by rearing Atlantic cod Gadus morhua L. larvae in 3 pCO2 concentrations-control (370 µatm), medium (1800 µatm) and high (4200 µatm)-from March to May 2010. Increased otolith growth was observed in 7 to 46 d post hatch (dph) cod larvae at elevated pCO2 concentrations. The sagittae and lapilli were usually largest in the high pCO2 treatment followed by the medium and control treatments. The greatest difference in mean otolith surface area (normalized to fish length) was for sagittae at 11 dph, with medium and high treatments being 46 and 43% larger than the control group, respectively. There was no significant pCO2 effect on the shape of the otoliths nor were there any trends in the fluctuating asymmetry, defined as the difference between the right and left sides, in relation to the increase in otolith growth from elevated pCO2.

Seawater carbonate chemistry, calcification and photosynthesis during experiments with corals, 2005

An investigation was conducted to determine the effects of elevated pCO2 on the net production and calcification of an assemblage of corals maintained under near-natural conditions of temperature, light, nutrient, and flow. Experiments were performed in summer and winter to explore possible interactions between seasonal change in temperature and irradiance and the effect of elevated pCO2. Particular attention was paid to interactions between net production and calcification because these two processes are thought to compete for the same internal supply of dissolved inorganic carbon (DIC). A nutrient enrichment experiment was performed because it has been shown to induce a competitive interaction between photosynthesis and calcification that may serve as an analog to the effect of elevated pCO2. Net carbon production, NPC, increased with increased pCO2 at the rate of 3 ± 2% (?mol CO2aq kg?1)?1. Seasonal change of the slope NPC-[CO2aq] relationship was not significant. Calcification (G) was strongly related to the aragonite saturation state ? a . Seasonal change of the G-? a relationship was not significant. The first-order saturation state model gave a good fit to the pooled summer and winter data: G = (8 ± 1 mmol CaCO3 m?2 h?1)(? a ? 1), r 2 = 0.87, P = 0.0001. Both nutrient and CO2 enrichment resulted in an increase in NPC and a decrease in G, giving support to the hypothesis that the cellular mechanism underlying the decrease in calcification in response to increased pCO2 could be competition between photosynthesis and calcification for a limited supply of DIC.

Seawater carbonate chemistry and shell length of Mediterranean pteropod Cavolinia inflexa larvae during experiments

Larvae of the Mediterranean pteropod Cavolinia inflexa were maintained at controlled pHT values of 8.1, 7.82 and 7.51, equivalent respectively to pCO2 levels of 380, 857 and 1713 µatm. At pHT 7.82 larvae exhibited malformations and lower shell growth, compared to the control condition. At pHT 7.51 the larvae did not make shells but were viable and showed a normal development. However, smaller shells or no shells will have both ecological (food web) and biogeochemical (export of carbon and carbonate) consequences. These results confirm that 1pteropods, as well as the species dependent upon them as a food resource, will be severely impacted by ocean acidification.

Seawater carbonate chemistry and processes during experiments with benthic foraminifera Ammonia tepida

Evidence of increasing concentrations of dissolved carbon dioxide, especially in the surface ocean and its associated impacts on calcifying organisms, is accumulating. Among these organisms, benthic and planktonic foraminifera are responsible for a large amount of the globally precipitated calcium carbonate. Hence, their response to an acidifying ocean may have important consequences for future inorganic carbon cycling. To assess the sensitivity of benthic foraminifera to changing carbon dioxide levels and subsequent alteration in seawater carbonate chemistry, we cultured specimens of the shallow water species Ammonia tepida at two concentrations of atmospheric CO2 (230 and 1900 ppmv) and two temperatures (10 °C and 15 °C). Shell weights and elemental compositions were determined. Impact of high and low pCO2 on elemental composition are compared with results of a previous experiment were specimens were grown under ambient conditions (380 ppvm, no shell weight measurements of specimen grown under ambient conditions are, however, available). Results indicate that shell weights decrease with decreasing [CO3], although calcification was observed even in the presence of calcium carbonate under-saturation, and also decrease with increasing temperature. Thus both warming and ocean acidification may act to decrease shell weights in the future. Changes in [CO3] or total dissolved inorganic carbon do not affect the Mg distribution coefficient. On the contrary, Sr incorporation is enhanced under increasing [CO3]. Implications of these results for the paleoceanographic application of foraminifera are discussed.

Seawater carbonate chemistry and calcification rate during mesocosm experiments with coral Montipora capitata, 2008

A long-term (10 months) controlled experiment was conducted to test the impact of increased partial pressure of carbon dioxide (pCO2) on common calcifying coral reef organisms. The experiment was conducted in replicate continuous flow coral reef mesocosms flushed with unfiltered sea water from Kaneohe Bay, Oahu, Hawaii. Mesocosms were located in full sunlight and experienced diurnal and seasonal fluctuations in temperature and sea water chemistry characteristic of the adjacent reef flat. Treatment mesocosms were manipulated to simulate an increase in pCO2 to levels expected in this century [midday pCO2 levels exceeding control mesocosms by 365 ± 130 µatm (mean ± sd)]. Acidification had a profound impact on the development and growth of crustose coralline algae (CCA) populations. During the experiment, CCA developed 25% cover in the control mesocosms and only 4% in the acidified mesocosms, representing an 86% relative reduction. Free-living associations of CCA known as rhodoliths living in the control mesocosms grew at a rate of 0.6 g buoyant weight per year while those in the acidified experimental treatment decreased in weight at a rate of 0.9 g buoyant weight per year, representing a 250% difference. CCA play an important role in the growth and stabilization of carbonate reefs, so future changes of this magnitude could greatly impact coral reefs throughout the world. Coral calcification decreased between 15% and 20% under acidified conditions. Linear extension decreased by 14% under acidified conditions in one experiment. Larvae of the coral Pocillopora damicornis were able to recruit under the acidified conditions. In addition, there was no significant difference in production of gametes by the coral Montipora capitata after 6 months of exposure to the treatments.

Seawater carbonate chemistry, growth rate and Emiliania huxleyi (strain AC481) biological processes during experiments, 2010

The impact of ocean acidification and increased water temperature on marine ecosystems, in particular those involving calcifying organisms, has been gradually recognised. We examined the individual and combined effects of increased pCO2 (180 ppmV CO2, 380 ppmV CO2 and 750 ppmV CO2 corresponding to past, present and future CO2 conditions, respectively) and temperature (13 °C and 18 °C) during the exponential growth phase of the coccolithophore E. huxleyi using batch culture experiments. We showed that cellular production rate of Particulate Organic Carbon (POC) increased from the present to the future CO2 treatments at 13 °C. A significant effect of pCO2 and of temperature on calcification was found, manifesting itself in a lower cellular production rate of Particulate Inorganic Carbon (PIC) as well as a lower PIC:POC ratio at future CO2 levels and at 18 °C. Coccosphere-sized particles showed a size reduction with both increasing temperature and CO2concentration. The influence of the different treatments on coccolith morphology was studied by categorizing SEM coccolith micrographs. The number of well-formed coccoliths decreased with increasing pCO2 while temperature did not have a significant impact on coccolith morphology. No interacting effects of pCO2 and temperature were observed on calcite production, coccolith morphology or on coccosphere size. Finally, our results suggest that ocean acidification might have a larger adverse impact on coccolithophorid calcification than surface water warming.