Seawater carbonate chemistry and buffer capacity of the coelomic fluid in echinoderms in a laboratory experiment

The increase in atmospheric CO2 due to anthropogenic activity results in an acidification of the surface waters of the oceans. The impact of these chemical changes depends on the considered organisms. In particular, it depends on the ability of the organism to control the pH of its inner fluids. Among echinoderms, this ability seems to differ significantly according to species or taxa. In the present paper, we investigated the buffer capacity of the coelomic fluid in different echinoderm taxa as well as factors modifying this capacity. Euechinoidea (sea urchins except Cidaroidea) present a very high buffer capacity of the coelomic fluid (from 0.8 to 1.8 mmol/kg SW above that of seawater), while Cidaroidea (other sea urchins), starfish and holothurians have a significantly lower one (from -0.1 to 0.4 mmol/kg SW compared to seawater). We hypothesize that this is linked to the more efficient gas exchange structures present in the three last taxa, whereas Euechinoidea evolved specific buffer systems to compensate lower gas exchange abilities. The constituents of the buffer capacity and the factors influencing it were investigated in the sea urchin Paracentrotus lividus and the starfish Asterias rubens. Buffer capacity is primarily due to the bicarbonate buffer system of seawater (representing about 63% for sea urchins and 92% for starfish). It is also partly due to coelomocytes present in the coelomic fluid (around 8% for both) and, in P. lividus only, a compound of an apparent size larger than 3 kDa is involved (about 15%). Feeding increased the buffer capacity in P. lividus (to a difference with seawater of about 2.3 mmol/kg SW compared to unfed ones who showed a difference of about 0.5 mmol/kg SW) but not in A. rubens (difference with seawater of about 0.2 for both conditions). In P. lividus, decreased seawater pH induced an increase of the buffer capacity of individuals maintained at pH 7.7 to about twice that of the control individuals and, for those at pH 7.4, about three times. This allowed a partial compensation of the coelomic fluid pH for individuals maintained at pH 7.7 but not for those at pH 7.4.

Euechinoidea and Cidaroidea respond differently to ocean acidification

The impact of the chemical changes in the ocean waters due to the increasing atmospheric CO2 depends on the ability of an organism to control extracellular pH. Among sea urchins, this seems specific to the Euechinoidea, sea urchins except Cidaroidea. However, Cidaroidea survived two ocean acidification periods: the Permian-Trias and the Cretaceous-Tertiary crises. We investigated the response of these two sea urchin groups to reduced seawater pH with the tropical cidaroid Eucidaris tribuloides, the sympatric euechinoid Tripneustes ventricosus and the temperate euechinoid Paracentrotus lividus. Both euechinoid showed a compensation of the coelomic fluid pH due to increased buffer capacity. This was linked to an increased concentration of DIC in the coelomic fluid and thus of bicarbonate ions (most probably originating from the surrounding seawater as isotopic signature of the carbon -delta 13C- was similar). On the other hand, the cidaroid showed no changes within the coelomic fluid. Moreover, the delta 13C of the coelomic fluid did not match that of the seawater and was not significantly different between the urchins from the different treatments. Feeding rate was not affected in any species. While euechinoids are able to regulate their extracellular acid-base balance, many questions are still unanswered on the costs of this capacity. On the contrary, cidaroids do not seem affected by a reduced seawater pH. Further investigations need to be undertaken to cover more species and physiological and metabolic parameters in order to determine if energy trade-offs occur and how this mechanism of compensation is distributed among sea urchins.

Seawater carbonate chemistry and calcium carbonate of Padina spp., photosynthesis of Padina pavonica in nature CO2 gradients experiment

Predicting the impacts of ocean acidification on coastal ecosystems requires an understanding of the effects on macroalgae and their grazers, as these underpin the ecology of rocky shores. Whilst calcified coralline algae (Rhodophyta) appear to be especially vulnerable to ocean acidification, there is a lack of information concerning calcified brown algae (Phaeophyta), which are not obligate calcifiers but are still important producers of calcium carbonate and organic matter in shallow coastal waters. Here, we compare ecological shifts in subtidal rocky shore systems along CO2 gradients created by volcanic seeps in the Mediterranean and Papua New Guinea, focussing on abundant macroalgae and grazing sea urchins. In both the temperate and tropical systems the abundances of grazing sea urchins declined dramatically along CO2 gradients. Temperate and tropical species of the calcifying macroalgal genus Padina (Dictyoaceae, Phaeophyta) showed reductions in CaCO3 content with CO2 enrichment. In contrast to other studies of calcified macroalgae, however, we observed an increase in the abundance of Padina spp. in acidified conditions. Reduced sea urchin grazing pressure and significant increases in photosynthetic rates may explain the unexpected success of decalcified Padina spp. at elevated levels of CO2. This is the first study to provide a comparison of ecological changes along CO2 gradients between temperate and tropical rocky shores. The similarities we found in the responses of Padina spp. and sea urchin abundance at several vent systems increases confidence in predictions of the ecological impacts of ocean acidification over a large geographical range.

Seawater carbonate chemistry and Strongylocentrotus droebachiensis larval and juvenile survival and reporductive processes, 2012

Anthropogenic CO2 emissions are acidifying the world's oceans. A growing body of evidence demonstrates that ocean acidification can impact survival, growth, development and physiology of marine invertebrates. Here we tested the impact of long term (up to 16 months) and trans life-cycle (adult, embryo/larvae and juvenile) exposure to elevated pCO2 (1200 µatm, compared to control 400 µatm) on the green sea urchin Strongylocentrotus droebachiensis. Female fecundity was decreased 4.5 fold when acclimated to elevated pCO2 for 4 months during reproductive conditioning while no difference was observed in females acclimated for 16 months. Moreover, adult pre-exposure for 4 months to elevated pCO2, had a direct negative impact on subsequent larval settlement success. Five to nine times fewer offspring reached the juvenile stage in cultures using gametes collected from adults previously acclimated to high pCO2 for 4 months. However, no difference in larval survival was observed when adults were pre-exposed for 16 months to elevated pCO2. pCO2 had no direct negative impact on juvenile survival except when both larvae and juveniles were raised in elevated pCO2. These negative effects on settlement success and juvenile survival can be attributed to carry-over effects from adults to larvae and from larvae to juveniles. Our results support the contention that adult sea urchins can acclimate to moderately elevated pCO2 in a matter of a few months and that carry-over effects can exacerbate the negative impact of ocean acidification on larvae and juveniles.

Seawater carbonate chemistry and calcification of the crustose coralline alga Hydrolithon onkodes in a laboratory experiment

Anthropogenic CO2 emissions have exacerbated two environmental stressors, global climate warming and ocean acidification (OA), that have serious implications for marine ecosystems. Coral reefs are vulnerable to climate change yet few studies have explored the potential for interactive effects of warming temperature and OA on an important coral reef calcifier, crustose coralline algae (CCA). Coralline algae serve many important ecosystem functions on coral reefs and are one of the most sensitive organisms to ocean acidification. We investigated the effects of elevated pCO2 and temperature on calcification of Hydrolithon onkodes, an important species of reef-building coralline algae, and the subsequent effects on susceptibility to grazing by sea urchins. H. onkodes was exposed to a fully factorial combination of pCO2 (420, 530, 830 µatm) and temperature (26, 29 °C) treatments, and calcification was measured by the change in buoyant weight after 21 days of treatment exposure. Temperature and pCO2 had a significant interactive effect on net calcification of H. onkodes that was driven by the increased calcification response to moderately elevated pCO2. We demonstrate that the CCA calcification response was variable and non-linear, and that there was a trend for highest calcification at ambient temperature. H. onkodes then was exposed to grazing by the sea urchin Echinothrix diadema, and grazing was quantified by the change in CCA buoyant weight from grazing trials. E. diadema removed 60% more CaCO3 from H. onkodes grown at high temperature and high pCO2 than at ambient temperature and low pCO2. The increased susceptibility to grazing in the high pCO2 treatment is among the first evidence indicating the potential for cascading effects of OA and temperature on coral reef organisms and their ecological interactions.

CO2-induced fertilization impairment in Strongylocentrotus droebachiensis collected in the Arctic

Fertilization depends on distribution and aggregation patterns of sea urchins which influence gamete contact time and may potentially enhance their vulnerability to ocean acidification. In this study, we conducted fertilization experiments to assess the effects of selected pH scenarios on fertilization success of Strongylocentrotus droebachiensis, from Spitsbergen, Arctic. Acidification was achieved by aerating seawater with different CO2 partial pressures to represent pre-industrial and present conditions (measured ~180-425 µatm) and future acidification scenarios (~550-800, ~1,300, ~2,000 µatm). Fertilization success was defined as the proportion of successful/unsuccessful fertilizations per treatment; eggs were classified according to features of their fertilization envelope (FE), hyaline layer (HL) and achievement of cellular division. The diagnostic findings of specific pathological aberrations were described in detail. We additionally measured intracellular pH changes in unfertilized eggs exposed for 1 h to selected acidification treatments using BCECF/AM. We conclude that (a) acidified conditions increase the proportion of eggs that failed fertilization, (b) acidification may increase the risk of polyspermy due to failures in the FE formation supported by the occasional observation of multiple sperms in the perivitelline space and (c) irregular formation of the embryo may arise due to impaired formation of the HL. The decrease in fertilization success could be also related to the observed changes in intracellular pH at pCO2 ~ 1,000 µatm or higher.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.

Seawater carbonate chemistry and processes during experiments with seaurchins Hemicentrotus pulcherrimus and Echinometra mathaei, 2004

Increased carbon dioxide (CO2) concentration in the atmosphere will change the balance of the components of carbonate chemistry and reduce the pH at the ocean surface. Here, we report the effects of increased CO2 concentration on the early development of the sea urchins Hemicentrotus pulcherrimus and Echinometra mathaei. We examined the fertilization, early cleavage, and pluteus larval stage to evaluate the impact of elevated CO2 concentration on fertilization rate, cleavage rate, developmental speed, and pluteus larval morphology. Furthermore, we compared the effects of CO2 and HCl at the same pH in an attempt to elucidate any differences between the two. We found that fertilization rate, cleavage rate, developmental speed, and pluteus larval size all tended to decrease with increasing CO2 concentration. Furthermore, CO2-seawater had a more severe effect than HCl-seawater on the fertilization rate. By contrast, the effects on cleavage rate, developmental speed, and pluteus larval morphology were similar for CO2- and HCl-seawater. Our results suggest that both decreased pH and altered carbonate chemistry affect the early development and life history of marine animals, implying that increased seawater CO2 concentration will seriously alter marine ecosystems. The effects of CO2 itself on marine organisms therefore requires further clarification.

Stable isotope records of corals from the northern Red Sea

We present a study based on X-ray chronologies and the stable isotopic composition of fossil Porites spp. corals from the northern Gulf of Aqaba (Red Sea) covering the mid-Holocene period from 5750 to 4450 14C years BP (before present). The stable oxygen and carbon isotopic compositions of five specimens reveal regular annual periodicities. Compared with modern Porites spp. from the same environment, the average seasonal delta18O amplitude of the fossil corals is higher (by ca. 0.35-0.60?), whereas annual growth rates are lower (by ca. 3.5 to 2 mm/year). This suggests stronger seasonality of sea surface temperatures and increased variability of the oxygen isotopic composition of the sea water due to changes in the precipitation and evaporation regime during the mid-Holocene. Most likely, summer monsoon rains reached the northern end of the Red Sea at that time. Average annual coral growth rates are diminished probably due to an increased input and resuspension of terrestrial debris to the shallow marine environment during more humid conditions. Our results corroborate published reports of paleodata and model simulations suggesting a northward migration of the African monsoon giving rise to increased seasonalities during the mid-Holocene over northeastern Africa and Arabia.

Investigation of Pliocene foraminifera on sediment core CRP-1 from the Ross Sea, Antarctica

Mixed assemblages of Pliocene and Quaternary foraminifera occur within the Quaternary succession of the CRP-1 drillhole. Pliocene foraminifera are not present in the lowermost Unit 4.1. are rare in Unit 3.1 and 2.3, are relatively common in Units 2.2 and 2.1, and are absent in Unit 1.1. Fifteen and twelve species were documented in two of the samples from Units 2.2 and 2.1 respectively. A census count of foraminifera in a sample at 26.89 mbsf (Unit 2.2) indicated that 39% of the tests were from a Pliocene source, with the remaining 61% tests assigned to the in situ Quaternary assemblage. There appears to be a close correlation between the stratigraphic distribution of ice-rafted sediments and the test number and diversity of Pliocene taxa. It is concluded that Pliocene assemblages were not derived from submarine outcrops on Roberts Ridge, but are more likely to have been rafted to the site via major trunk valley drainage systems such as operated within the Mackay and Ferrar glacial valleys. The co-occurrence of marine biota (including foraminifera), fossil wood, pollen, and igneous clasts in the Quaternary succession of CRP-l, points to the marine and terrestrial facies of the Pliocene Sirius Group as a likely source. A major episode of erosion and transport of sediment into the offshore marine basins at about ~1 Ma may have been triggered by dynamism in the ice sheet-glacier system, an episode of regional uplift in the Transantarctic Mountains, sea level oscillations and associated changes in the land-to-sea drainage baselines, or some combination of these factors.

Planktonic foraminifera fluxes and stable isotope ratios in the Irminger Sea

Past water column stratification can be assessed through comparison of the d18O of different planktonic foraminiferal species. The underlying assumption is that different species form their shells simultaneously, but at different depths in the water column. We evaluate this assumption using a sediment trap time-series of Neogloboquadrina pachyderma (s) and Globigerina bulloides from the NW North Atlantic. We determined fluxes, d18O and d13C of shells from two size fractions to assess size-related effects on shell chemistry and to better constrain the underlying causes of isotopic differences between foraminifera in deep-sea sediments. Our data indicate that in the subpolar North Atlantic differences in the seasonality of the shell flux, and not in depth habitat or test size, determine the interspecies Delta d18O. N. pachyderma (s) preferentially forms from early spring to late summer, whereas the flux ofG. bulloides peaks later in the season and is sustained until autumn. Likewise, seasonality influences large and small specimens differently, with large shells settling earlier in the season. The similarity of the seasonal d18O patterns between the two species indicates that they calcify in an overlapping depth zone close to the surface. However, their d13C patterns are markedly different (>1 per mil). Both species have a seasonally variable offset from d13CDIC that appears to be governed primarily by temperature, with larger offsets associated with higher temperatures. The variable offset from d13CDIC implies that seasonality of the flux affects the fossil d13C signal, which has implications for reconstruction of the past oceanic carbon cycle.