Seawater carbonate chemistry and community calcification during Miyako Island (Japan) coral reef studies, 1994

Coral reefs are characterized by enormous carbonate production of the organisms. It is known that rapid calcification is linked to photosynthesis under control of the carbonate equilibrium in seawater. We have established a model simulating the coexisting states of photosynthesis and calcification in order to examine the effects of photosynthesis and calcification on the carbonate system in seawater. Supposing that the rates of photosynthesis and calcification are proportional to concentrations of their inorganic carbon source, the model calculations indicate that three kinds of unique interactions of the organic and inorganic carbon productions are expected. These are photosynthetic enhancement of calcification, calcification which benefits photosynthesis and carbonate dissolution induced by respiration. The first effect appears when the photosynthetic rate is more than approximately 1.2 larger than that of calcification. This effect is caused by the increase of CO3 content and carbonate saturation degree in seawater. If photosynthesis use molecular carbon dioxide, the second effect occurs when the calcification rate is more than approximately 1.6 times larger than that of photosynthesis. Time series model experiments indicate that photosynthesis and calcification potentially enhance each other and that organic and inorganic carbon is produced more efficiently in the coexisting system than in the isolated reactions. These coexisting effects on production enhancement of photosynthesis and calcification are expected to appear not only in the internal pool of organisms but also in a reef environment which is isolated from the outer ocean during low tide. According to the measurements on the fringing type Shiraho Reef in the Ryukyu Islands, the diurnal change of water properties (pH, total alkalinity, total carbon dioxide and carbonate saturation degree) were conspicuous. This environment offers an appropriate condition for the appearance of these coexisting effects. The photosynthetic enhancement of calcification and the respiratory inducement of decalcification were observed during day-time and night-time slack-water periods, respectively. These coexisting effects, especially the photosynthetic enhancement of calcification, appear to play important roles for fluorishing coral reef communities.

Limited effects of increased CO2 and temperature on metal and radionuclide bioaccumulation in a sessile invertebrate, the oyster Crassostrea gigas

This study investigated the combined effects of reduced pH and increased temperature on the capacities of the Pacific cupped oyster Crassostrea gigas to bioconcentrate radionuclide and metals. Oysters were exposed to dissolved radiotracers (110mAg, 241Am, 109Cd,57Co,54Mn, and 65Zn) at three pH (7.5, 7.8, 8.1) and two temperatures (21 and 24°C) under controlled laboratory conditions. Although calcifying organisms are recognized as particularly vulnerable to ocean acidification, the oyster did not accumulate differently the studied metals when exposed under the different pH conditions. However, temperature alone or in combination with pH somewhat altered the bioaccumulation of the studied elements. At pH 7.5, Cd was accumulated with an uptake rate constant twofold higher at 24°C than 21°C. Bioaccumulation of Mn was significantly affected by an interactive effect between seawater pH and temperature, with a decreased uptake rate at pH 7.5 when temperature increased (27 ± 1 vs. 17 ± 1 /day at 21 and 24°C, respectively). Retention of Co and Mn tended also to decrease at the same pH with decreasing temperature. Neither pH nor temperature affected strongly the elements distribution between shell and soft tissues. Significant effects of pH were found on the bioaccessibility of Mn, Zn, and 241Am during experimental in vitro simulation of human digestion.

Laser ablation data of benthic foraminifera from surface sediments collected from the muddy intertidal flats near Dorum, Northwestern Germany in Autumn 2008

Biological activity introduces variability in element incorporation during calcification and thereby decreases the precision and accuracy when using foraminifera as geochemical proxies in paleoceanography. This so-called 'vital effect' consists of organismal and environmental components. Whereas organismal effects include uptake of ions from seawater and subsequent processing upon calcification, environmental effects include migration- and seasonality-induced differences. Triggering asexual reproduction and culturing juveniles of the benthic foraminifer Ammonia tepida under constant, controlled conditions allow environmental and genetic variability to be removed and the effect of cell-physiological controls on element incorporation to be quantified. Three groups of clones were cultured under constant conditions while determining their growth rates, size-normalized weights and single-chamber Mg/Ca and Sr/Ca using laser ablation-inductively coupled plasma-mass spectrometry (LA-ICP-MS). Results show no detectable ontogenetic control on the incorporation of these elements in the species studied here. Despite constant culturing conditions, Mg/Ca varies by a factor of similar to 4 within an individual foraminifer while intra-individual Sr/Ca varies by only a factor of 1.6. Differences between clone groups were similar to the intra-clone group variability in element composition, suggesting that any genetic differences between the clone-groups studied here do not affect trace element partitioning. Instead, variability in Mg/Ca appears to be inherent to the process of bio-calcification itself. The variability in Mg/Ca between chambers shows that measurements of at least 6 different chambers are required to determine the mean Mg/Ca value for a cultured foraminiferal test with a precision of <= 10%

Geographical variation in shell morphology of juvenile snails (Concholepas concholepas) along the physical-chemical gradient of the Chilean coast

Changes in phenotypic traits, such as mollusc shells, are indicative of variations in selective pressure along environmental gradients. Recently, increased sea surface temperature (SST) and ocean acidification (OA) due to increased levels of carbon dioxide in the seawater have been described as selective agents that may affect the biological processes underlying shell formation in calcifying marine organisms. The benthic snail Concholepas concholepas (Muricidae) is widely distributed along the Chilean coast, and so is naturally exposed to a strong physical-chemical latitudinal gradient. In this study, based on elliptical Fourier analysis, we assess changes in shell morphology (outlines analysis) in juvenile C. concholepas collected at northern (23°S), central (33°S) and southern (39°S) locations off the Chilean coast. Shell morphology of individuals collected in northern and central regions correspond to extreme morphotypes, which is in agreement with both the observed regional differences in the shell apex outlines, the high reclassification success of individuals (discriminant function analysis) collected in these regions, and the scaling relationship in shell weight variability among regions. However, these extreme morphotypes showed similar patterns of mineralization of calcium carbonate forms (calcite and aragonite). Geographical variability in shell shape of C. concholepas described by discriminant functions was partially explained by environmental variables (pCO2, SST). This suggests the influence of corrosive waters, such as upwelling and freshwaters penetrating into the coastal ocean, upon spatial variation in shell morphology. Changes in the proportion of calcium carbonate forms precipitated by C. concholepas across their shells and its susceptibility to corrosive coastal waters are discussed.

Decline in Coccolithophore Diversity and Impact on Coccolith Morphogenesis Along a Natural CO2 Gradient

A natural pH gradient caused by marine CO2 seeps off Vulcano Island (Italy) was used to assess the effects of ocean acidification on coccolithophores, which are abundant planktonic unicellular calcifiers. Such seeps are used as natural laboratories to study the effects of ocean acidification on marine ecosystems, since they cause long-term changes in seawater carbonate chemistry and pH, exposing the organisms to elevated CO2 concentrations and therefore mimicking future scenarios. Previous work at CO2 seeps has focused exclusively on benthic organisms. Here we show progressive depletion of 27 coccolithophore species, in terms of cell concentrations and diversity, along a calcite saturation gradient from Omega calcite 6.4 to <1. Water collected close to the main CO2 seeps had the highest concentrations of malformed Emiliania huxleyi. These observations add to a growing body of evidence that ocean acidification may benefit some algae but will likely cause marine biodiversity loss, especially by impacting calcifying species, which are affected as carbonate saturation falls.

Seawater carbonate chemistry and larval development and settlement of the sea urchin Arbacia lixula in a laboratory experiment

We studied the effects of temperature and pH on larval development, settlement and juvenile survival of a Mediterranean population of the sea urchin Arbacia lixula. Three temperatures (16, 17.5 and 19 °C) were tested at present pH conditions (pHT 8.1). At 19 °C, two pH levels were compared to reflect present average (pHT 8.1) and near-future average conditions (pHT 7.7, expected by 2100). Larvae were reared for 52-days to achieve the full larval development and complete the metamorphosis to the settler stage. We analyzed larval survival, growth, morphology and settlement success. We also tested the carry-over effect of acidification on juvenile survival after 3 days. Our results showed that larval survival and size significantly increased with temperature. Acidification resulted in higher survival rates and developmental delay. Larval morphology was significantly altered by low temperatures, which led to narrower larvae with relatively shorter skeletal rods, but larval morphology was only marginally affected by acidification. No carry-over effects between larvae and juveniles were detected in early settler survival, though settlers from larvae reared at pH 7.7 were significantly smaller than their counterparts developed at pH 8.1. These results suggest an overall positive effect of environmental parameters related to global change on the reproduction of A. lixula, and reinforce the concerns about the increasing negative impact on shallow Mediterranean ecosystems of this post-glacial colonizer.

Seawater carbonate chemistry and resource allocation and extracellular acid-base status in the sea urchin Strongylocentrotus droebachiensis during experiments, 2012

Anthropogenic CO2 emission will lead to an increase in seawater pCO2 of up to 80-100 Pa (800-1000 µatm) within this century and to an acidification of the oceans. Green sea urchins (Strongylocentrotus droebachiensis) occurring in Kattegat experience seasonal hypercapnic and hypoxic conditions already today. Thus, anthropogenic CO2 emissions will add up to existing values and will lead to even higher pCO2 values >200 Pa (>2000 µatm). To estimate the green sea urchins' potential to acclimate to acidified seawater, we calculated an energy budget and determined the extracellular acid base status of adult S. droebachiensis exposed to moderately (102 to 145 Pa, 1007 to 1431 µatm) and highly (284 to 385 Pa, 2800 to 3800 µatm) elevated seawater pCO2 for 10 and 45 days. A 45 - day exposure to elevated pCO2 resulted in a shift in energy budgets, leading to reduced somatic and reproductive growth. Metabolic rates were not significantly affected, but ammonium excretion increased in response to elevated pCO2. This led to decreased O:N ratios. These findings suggest that protein metabolism is possibly enhanced under elevated pCO2 in order to support ion homeostasis by increasing net acid extrusion. The perivisceral coelomic fluid acid-base status revealed that S. droebachiensis is able to fully (intermediate pCO2) or partially (high pCO2) compensate extracellular pH (pHe) changes by accumulation of bicarbonate (maximum increases 2.5 mM), albeit at a slower rate than typically observed in other taxa (10 day duration for full pHe compensation). At intermediate pCO2, sea urchins were able to maintain fully compensated pHe for 45 days. Sea urchins from the higher pCO2 treatment could be divided into two groups following medium-term acclimation: one group of experimental animals (29%) contained remnants of food in their digestive system and maintained partially compensated pHe (+2.3 mM HCO3), while the other group (71%) exhibited an empty digestive system and a severe metabolic acidosis (-0.5 pH units, -2.4 mM HCO3). There was no difference in mortality between the three pCO2 treatments. The results of this study suggest that S. droebachiensis occurring in the Kattegat might be pre-adapted to hypercapnia due to natural variability in pCO2 in its habitat. We show for the first time that some echinoderm species can actively compensate extracellular pH. Seawater pCO2 values of >200 Pa, which will occur in the Kattegat within this century during seasonal hypoxic events, can possibly only be endured for a short time period of a few weeks. Increases in anthropogenic CO2 emissions and leakages from potential sub-seabed CO2 storage (CCS) sites thus impose a threat to the ecologically and economically important species S. droebachiensis.

Acid-base regulation in the Dungeness crab (Metacarcinus magister)

Homeostatic regulation allows organisms to secure basic physiological processes in a varying environment. To counteract fluctuations in ambient carbonate system speciation due to elevated seawater pCO2 (hypercapnia), many aquatic crustaceans excrete/accumulate acid-base equivalents through their gills; however, not much is known about the role of ammonia in this response. The present study investigated the effects of hypercapnia on acid-base and ammonia regulation in the Dungeness crab, Metacarcinus magister on the whole animal and isolated gill levels. Hemolymph pCO2 and [HCO3]- increased in M. magister acclimated to elevated pCO2 (330 Pa), while pH remained stable. Additionally, hemolymph [Na+], [Ca2+], and [SO4]2- were significantly increased. When challenged with varying pH during gill perfusion, the pH of the artificial hemolymph remained relatively unchanged. Overall, ammonia production and excretion, as well as oxygen consumption, were reduced in crabs acclimated to elevated pCO2, demonstrating that either (amino acid) oxidation is reduced in response to this particular stress, or nitrogenous wastes are excreted in an alternative form.

Impacts of elevated CO2 on particulate and dissolved organic matter production: microcosm experiments using iron-deficient plankton communities in open subarctic waters

Response of phytoplankton to increasing CO2 in seawater in terms of physiology and ecology is key to predicting changes in marine ecosystems. However, responses of natural plankton communities especially in the open ocean to higher CO2 levels have not been fully examined. We conducted CO2 manipulation experiments in the Bering Sea and the central subarctic Pacific, known as high nutrient and low chlorophyll regions, in summer 2007 to investigate the response of organic matter production in iron-deficient plankton communities to CO2 increases. During the 14-day incubations of surface waters with natural plankton assemblages in microcosms under multiple pCO2 levels, the dynamics of particulate organic carbon (POC) and nitrogen (PN), and dissolved organic carbon (DOC) and phosphorus (DOP) were examined with the plankton community compositions. In the Bering site, net production of POC, PN, and DOP relative to net chlorophyll-a production decreased with increasing pCO2. While net produced POC:PN did not show any CO2-related variations, net produced DOC:DOP increased with increasing pCO2. On the other hand, no apparent trends for these parameters were observed in the Pacific site. The contrasting results observed were probably due to the different plankton community compositions between the two sites, with plankton biomass dominated by large-sized diatoms in the Bering Sea versus ultra-eukaryotes in the Pacific Ocean. We conclude that the quantity and quality of the production of particulate and dissolved organic matter may be altered under future elevated CO2 environments in some iron-deficient ecosystems, while the impacts may be negligible in some systems.

Calcification rate of massive Porites spp. and Porites rus in the experiment of Moorea

This study tested the hypothesis that the response of corals to temperature and pCO2 is consistent between taxa. Juvenile massive Porites spp. and branches of P. rus from the back reef of Moorea were incubated for 1 month under combinations of temperature (29.3 °C and 25.6 °C) and pCO2 (41.6 Pa and 81.5 Pa) at an irradiance of 599 µmol quanta/m/s. Using microcosms and CO2 gas mixing technology, treatments were created in a partly nested design (tanks) with two between-plot factors (temperature and pCO2), and one within-plot factor (taxon); calcification was used as a dependent variable. pCO2 and temperature independently affected calcification, but the response differed between taxa. Massive Porites spp. was largely unaffected by the treatments, but P. rus grew 50% faster at 29.3 °C compared with 25.6 °C, and 28% slower at 81.5 Pa vs. 41.6 Pa CO2. A compilation of studies placed the present results in a broader context and tested the hypothesis that calcification for individual coral genera is independent of pH, [HCO3]-, and [CO3]2-. Unlike recent reviews, this analysis was restricted to studies reporting calcification in units that could be converted to nmol CaCO3/cm**2/h. The compilation revealed a high degree of variation in calcification as a function of pH, [HCO3]-, and [CO3]2-, and supported three conclusions: (1) studies of the effects of ocean acidification on corals need to pay closer attention to reducing variance in experimental outcomes to achieve stronger synthetic capacity, (2) coral genera respond in dissimilar ways to pH, [HCO3]-, and [CO3]2-, and (3) calcification of massive Porites spp. is relatively resistant to short exposures of increased pCO2, similar to that expected within 100 y.

Seawater carbonate chemistry and copepod Centropages tenuiremis feeding, filtering and respiration rate during experiments, 2012

Climate change mediates marine chemical and physical environments and therefore influences marine organisms. While increasing atmospheric CO2 level and associated ocean acidification has been predicted to stimulate marine primary productivity and may affect community structure, the processes that impact food chain and biological CO2 pump are less documented. We hypothesized that copepods, as the secondary marine producer, may respond to future changes in seawater carbonate chemistry associated with ocean acidification due to increasing atmospheric CO2 concentration. Here, we show that the copepod, Centropages tenuiremis, was able to perceive the chemical changes in seawater induced under elevated CO2 concentration (>1700 µatm, pH < 7.60) with avoidance strategy. The copepod's respiration increased at the elevated CO2 (1000 µatm), associated acidity (pH 7.83) and its feeding rates also increased correspondingly, except for the initial acclimating period, when it fed less. Our results imply that marine secondary producers increase their respiration and feeding rate in response to ocean acidification to balance the energy cost against increased acidity and CO2 concentration.