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The effects of dissolved inorganic carbon (DIC) on the growth of 3 red-tide dinoflagellates (Ceratium lineatum, Heterocapsa triquetra and Prorocentrum minimum) were studied at pH 8.0 and at higher pH levels, depending upon the pH tolerance of the individual species. The higher pH levels chosen for experiments were 8.55 for C. lineatum and 9.2 for the other 2 species. At pH 8.0, which approximates the pH found in the open sea, the maximum growth in all species was maintained until the total DIC concentration was reduced below ~0.4 and 0.2 mM for C. lineatum and the other 2 species, respectively. Growth compensation points (concentration of inorganic carbon needed for maintenance of cells) were reached at ~0.18 and 0.05 mM DIC for C. lineatum and the other 2 species, respectively. At higher pH levels, maximum growth rates were lower compared to growth at pH 8, even at very high DIC concentrations, indicating a direct pH effect on growth. Moreover, the concentration of bio-available inorganic carbon (CO2 + HCO3-) required for maintenance as well as the half-saturation constants were increased considerably at high pH compared to pH 8.0. Experiments with pH-drift were carried out at initial concentrations of 2.4 and 1.2 mM DIC to test whether pH or DIC was the main limiting factor at a natural range of DIC. Independent of the initial DIC concentrations, growth rates were similar in both incubations until pH had increased considerably. The results of this study demonstrated that growth of the 3 species was mainly limited by pH, while inorganic carbon limitation played a minor role only at very high pH levels and low initial DIC concentrations.

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Phytoplankton are the basis of marine food webs, and affect biogeochemical cycles. As CO2 levels increase, shifts in the frequencies and physiology of ecotypes within phytoplankton groups will affect their nutritional value and biogeochemical function. However, studies so far are based on a few representative genotypes from key species. Here, we measure changes in cellular function and growth rate at atmospheric CO2 concentrations predicted for the year 2100 in 16 ecotypes of the marine picoplankton Ostreococcus. We find that variation in plastic responses among ecotypes is on par with published between-genera variation, so the responses of one or a few ecotypes cannot estimate changes to the physiology or composition of a species under CO2 enrichment. We show that ecotypes best at taking advantage of CO2 enrichment by changing their photosynthesis rates most should increase in relative fitness, and so in frequency in a high-CO2 environment. Finally, information on sampling location, and not phylogenetic relatedness, is a good predictor of ecotypes likely to increase in frequency in this system.

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In recent years, some of the ramifications of the ocean acidification problematic derived from the anthropogenic rising of atmospheric CO2 have been widely studied. In particular, the potential effects of a lowering pH on tropical coral reefs have received special attention. However, only a few studies have focused on testing the effects of ocean acidification in corals from the Mediterranean Sea, despite the fact that this basin is especially sensitive to increasing atmospheric CO2. In this context, we investigated the response to ocean acidification of the two zooxanthellate coral species capable of constituting the main framework of the community, the endemic Cladocora caespitosa and the non-native Oculina patagonica. To this end, we examined the response of both species to pCO2 concentrations expected by the end of the century, 800 ppm, vs the present levels. Calcification rate measurements after 92 days of exposure to low pH conditions showed the same negative response in both species, a decrease of 32-35% compared to corals reared under control conditions. In addition, we detected in both species a correlation between the calcification rate of colonies in control conditions and the degree of impairment of the same colonies at low pH. Independent of species, faster growing colonies were more affected by decreased pH. After this period of decreased pH, we conducted a recovery experiment, in which corals reared in the acidic treatment were brought back to control conditions. In this case, normal calcification rates were reached in both species. Overall, our results suggest that O. patagonica and C. caespitosa will both be affected detrimentally by progressive ocean acidification in the near future. They do not display differences in response between native and non-native species but do manifest differential responses depending on calcification rate, pointing to a role of the coral genetics in determining the response of corals to ocean acidification.

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The effect of decreasing aragonite saturation state (Omega Arag) of seawater (elevated pCO2) on calcification rates of Acropora muricata was studied using nubbins prepared from parent colonies located at two sites of La Saline reef (La Réunion Island, western Indian Ocean): a back-reef site (BR) affected by nutrient-enriched groundwater discharge (mainly nitrate), and a reef flat site (RF) with low terrigenous inputs. Protein and chlorophyll a content of the nubbins, as well as zooxanthellae abundance, were lower at RF than BR. Nubbins were incubated at ~27°C over 2 h under sunlight, in filtered seawater manipulated to get differing initial pCO2 (1,440-340 µatm), Omega Arag (1.4-4.0), and dissolved inorganic carbon (DIC) concentrations (2,100-1,850 µmol kg-1). Increasing DIC concentrations at constant total alkalinity (AT) resulted in a decrease in Omega Arag and an increase in pCO2. AT at the beginning of the incubations was kept at a natural level of 2,193 +- 6 µmol kg-1 (mean +- SD). Net photosynthesis (NP) and calcification were calculated from changes in pH and AT during the incubations. Calcification decrease in response to doubling pCO2 relative to preindustrial level was 22% for RF nubbins. When normalized to surface area of the nubbins, (1) NP and calcification were higher at BR than RF, (2) NP increased in high pCO2 treatments at BR compared to low pCO2 treatments, and (3) calcification was not related to Omega Arag at BR. When normalized to NP, calcification was linearly related to Omega Arag at both sites, and the slopes of the relationships were not significantly different. The increase in NP at BR in the high pCO2 treatments may have increased calcification and thus masked the negative effect of low Omega Arag on calcification. Removing the effect of NP variations at BR showed that calcification declined in a similar manner with decreased Omega Arag (increased pCO2) whatever the nutrient loading.

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We investigated the effect of elevated partial pressure of CO2 (pCO2) on the photosynthesis and growth of four phylotypes (ITS2 types A1, A13, A2, and B1) from the genus Symbiodinium, a diverse dinoflagellate group that is important, both free-living and in symbiosis, for the viability of cnidarians and is thus a potentially important model dinoflagellate group. The response of Symbiodinium to an elevated pCO2 was phylotype-specific. Phylotypes A1 and B1 were largely unaffected by a doubling in pCO2 in contrast, the growth rate of A13 and the photosynthetic capacity of A2 both increased by ~ 60%. In no case was there an effect of ocean acidification (OA) upon respiration (dark- or light-dependent) for any of the phylotypes examined. Our observations suggest that OA might preferentially select among free-living populations of Symbiodinium, with implications for future symbioses that rely on algal acquisition from the environment (i.e., horizontal transmission). Furthermore, the carbon environment within the host could differentially affect the physiology of different Symbiodinium phylotypes. The range of responses we observed also highlights that the choice of species is an important consideration in OA research and that further investigation across phylogenetic diversity, for both the direction of effect and the underlying mechanism(s) involved, is warranted.

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The coccolithophore Calcidiscus leptoporus was grown in batch culture under nitrogen (N) as well as phosphorus (P) limitation. Growth rate, particulate inorganic carbon (PIC), particulate organic carbon (POC), particulate organic nitrogen (PON), and particulate organic phosphorus (POP) production were determined and coccolith morphology was analysed. While PON production decreased by 70% under N-limitation and POP production decreased by 65% under P-limitation, growth rate decreased by 33% under N- as well as P-limitation. POC as well as PIC production (calcification rate) increased by 27% relative to the control under P-limitation, and did not change under N-limitation. Coccolith morphology did not change in response to either P or N limitation. While these findings, supported by a literature survey, suggest that coccolith morphogenesis is not hampered by either P or N limitation, calcification rate might be. The latter conclusion is in apparent contradiction to our data. We discuss the reasons for this inference.

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The growth rate of Acropora cervicornis branch tips maintained in the laboratory was measured before, during, and after exposure to elevated nitrate (5 and 10 µM NO3-), phosphate (2 and 4 µM P-PO43) and/or pCO2 (CO2 ~700 to 800 µatm). The effect of increased pCO2 was greater than that of nutrient enrichment alone. High concentrations of nitrate or phosphate resulted in significant decreases in growth rate, in both the presence and absence of increased pCO2. The effect of nitrate and phosphate enrichment combined was additive or antagonistic relative to nutrient concentration and pCO2 level. Growth rate recovery was greater after exposure to increased nutrients or CO2 compared to increased nutrients and CO2. If these results accurately predict coral response in the natural environment, it is reasonable to speculate that the survival and reef-building potential of this species will be significantly negatively impacted by continued coastal nutrification and projected pCO2 increases.

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About 30% of the anthropogenically released CO2 is taken up by the oceans; such uptake causes surface ocean pH to decrease and is commonly referred to as ocean acidification (OA). Foraminifera are one of the most abundant groups of marine calcifiers, estimated to precipitate ca. 50 % of biogenic calcium carbonate in the open oceans. We have compiled the state of the art literature on OA effects on foraminifera, because the majority of OA research on this group was published within the last three years. Disparate responses of this important group of marine calcifiers to OA were reported, highlighting the importance of a process-based understanding of OA effects on foraminifera. We cultured the benthic foraminifer Ammonia sp. under a range of carbonate chemistry manipulation treatments to identify the parameter of the carbonate system causing the observed effects. This parameter identification is the first step towards a process-based understanding. We argue that CO3 is the parameter affecting foraminiferal size-normalized weights (SNWs) and growth rates. Based on the presented data, we can confirm the strong potential of Ammonia sp. foraminiferal SNW as a CO3 proxy.

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The Canary Basin lies in a region of strong interaction between the atmospheric and ocean circulation systems: Trade winds drive seasonal coastal upwelling and dust storm outbreaks from the neighbouring Sahara desert are the major source of terrigenous sediment. To investigate the forcing mechanisms for dust input and wind strength in the North Canary Basin, the temporal pattern of variability of sedimentological and geochemical proxy records has been analysed in two sediment cores between latitudes 30°30'N and 31°40'N. Spectral analysis of the dust proxy records indicates that insolation changes related to eccentricity and precession are the main periods of temporal variation in the record. Si/Al and grain-size of the terrigenous fraction show an increase in glacial-interglacial transitions while Al concentration and Fe/Al ratio are both in phase with minima in the precessional index. Hence, the results obtained show that the wind strength was intensified at Terminations. At times of maxima of Northern Hemisphere seasonal insolation, when the African monsoon was enhanced, the North Canary Basin also received higher dust input. This result suggests that the moisture brought by the monsoon may have increased the availability of dust in the source region.

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Anthropogenic carbon dioxide (CO2) emissions reduce pH of marine waters due to the absorption of atmospheric CO2 and formation of carbonic acid. Estuarine waters are more susceptible to acidification because they are subject to multiple acid sources and are less buffered than marine waters. Consequently, estuarine shell forming species may experience acidification sooner than marine species although the tolerance of estuarine calcifiers to pH changes is poorly understood. We analyzed 23 years of Chesapeake Bay water quality monitoring data and found that daytime average pH significantly decreased across polyhaline waters although pH has not significantly changed across mesohaline waters. In some tributaries that once supported large oyster populations, pH is increasing. Current average conditions within some tributaries however correspond to values that we found in laboratory studies to reduce oyster biocalcification rates or resulted in net shell dissolution. Calcification rates of juvenile eastern oysters, Crassostrea virginica, were measured in laboratory studies in a three-way factorial design with 3 pH levels, two salinities, and two temperatures. Biocalcification declined significantly with a reduction of ~0.5 pH units and higher temperature and salinity mitigated the decrease in biocalcification.

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Anthropogenic CO2 emissions are acidifying the world's oceans. A growing body of evidence is showing that ocean acidification impacts growth and developmental rates of marine invertebrates. Here we test the impact of elevated seawater pCO2 (129 Pa, 1271 µatm) on early development, larval metabolic and feeding rates in a marine model organism, the sea urchin Strongylocentrotus purpuratus. Growth and development was assessed by measuring total body length, body rod length, postoral rod length and posterolateral rod length. Comparing these parameters between treatments suggests that larvae suffer from a developmental delay (by ca. 8%) rather than from the previously postulated reductions in size at comparable developmental stages. Further, we found maximum increases in respiration rates of + 100 % under elevated pCO2, while body length corrected feeding rates did not differ between larvae from both treatments. Calculating scope for growth illustrates that larvae raised under high pCO2 spent an average of 39 to 45% of the available energy for somatic growth, while control larvae could allocate between 78 and 80% of the available energy into growth processes. Our results highlight the importance of defining a standard frame of reference when comparing a given parameter between treatments, as observed differences can be easily due to comparison of different larval ages with their specific set of biological characters.