Paleoceanography of sediment cores SO202-27-6 and MD01-2416

The glacial-to-Holocene evolution of subarctic Pacific surface water stratification and silicic acid (Si) dynamics is investigated based on new combined diatom oxygen (d18Odiat) and silicon (d30Sidiat) isotope records, along with new biogenic opal, subsurface foraminiferal d18O, alkenone-based sea surface temperature, sea ice, diatom, and core logging data from the NE Pacific. Our results suggest that d18Odiat values are primarily influenced by changes in freshwater discharge from the Cordilleran Ice Sheet (CIS), while corresponding d30Sidiat are primarily influenced by changes in Si supply to surface waters. Our data indicate enhanced glacial to mid Heinrich Stadial 1 (HS1) NE Pacific surface water stratification, generally limiting the Si supply to surface waters. However, we suggest that an increase in Si supply during early HS1, when surface waters were still stratified, is linked to increased North Pacific Intermediate Water formation. The coincidence between fresh surface waters during HS1 and enhanced ice-rafted debris sedimentation in the North Atlantic indicates a close link between CIS and Laurentide Ice Sheet dynamics and a dominant atmospheric control on CIS deglaciation. The Bølling/Allerød (B/A) is characterized by destratification in the subarctic Pacific and an increased supply of saline, Si-rich waters to surface waters. This change toward increased convection occurred prior to the Bølling warming and is likely triggered by a switch to sea ice-free conditions during late HS1. Our results furthermore indicate a decreased efficiency of the biological pump during late HS1 and the B/A (possibly also the Younger Dryas), suggesting that the subarctic Pacific has then been a source region of atmospheric CO2.

Growth, respiration and photophysiology of coral massive Porites spp. in the experiment of Moorea

I tested the hypothesis that high pCO2 (76.6 Pa and 87.2 Pa vs. 42.9 Pa) has no effect on the metabolism of juvenile massive Porites spp. after 11 days at 28 °C and 545 µmol quanta/m**2/s. The response was assessed as aerobic dark respiration, skeletal weight (i.e., calcification), biomass, and chlorophyll fluorescence. Corals were collected from the shallow (3-4 m) back reef of Moorea, French Polynesia (17°28.614'S, 149°48.917'W), and experiments conducted during April and May 2011. An increase in pCO2 to 76.6 Pa had no effect on any dependent variable, but 87.2 Pa pCO2 reduced area-normalized (but not biomass-normalized) respiration 36 %, as well as maximum photochemical efficiency (Fv/Fm) of open RCIIs and effective photochemical efficiency of RCIIs in actinic light (Delta F/F'm ); neither biomass, calcification, nor the energy expenditure coincident with calcification (J/g) was effected. These results do not support the hypothesis that high pCO2 reduces coral calcification through increased metabolic costs and, instead, suggest that high pCO2 causes metabolic depression and photochemical impairment similar to that associated with bleaching. Evidence of a pCO2 threshold between 76.6 and 87.2 Pa for inhibitory effects on respiration and photochemistry deserves further attention as it might signal the presence of unpredictable effects of rising pCO2.

Sterol, n-alcohol and n-alkane composition of living fen plants

In groundwater-fed fen peatlands, the surface biomass decays rapidly and, as a result, highly humified peat is formed. A high degree of humification constrains palaeoecological studies because reliable identification of plant remains is hampered. Organic geochemistry techniques as a means of identifying historical plant communities have been successfully applied tobog peat. The method has also been applied to fen peat, but without reference to the composition of fen plants. We have applied selected organic geochemistry methods to determine the composition of the neutral lipid fractions from 12 living fen plants, to investigate the potential for the distributions to characterize and separate different fen plants and plant groups. Our results show correspondence with previous studies, e.g. C23 and C25n-alkanes dominating Sphagnum spp. and C27 to C31 alkanes dominating vascular plants. However, we also found similarities in n-alkane distributions between Sphagnum spp. and the below ground parts of some vascular plants. We tested the efficiency of different n-alkane ratios to separate species and plant groups. The ratios used for bog studies (e.g. n-C23/n-C25 and n-C23/n-C29) did not work as consistently for fen plants. Some differences in sterol distribution were found between vascular plants and mosses; in general vascular plants had a higher concentration of sterols. When distributions of n-alkanes, n-alkane ratios and sterols were all included as variables, redundancy analysis (RDA) separated different plant groups into their own clusters. Our results imply that the pattern for bog biomarkers cannot directly be applied to fen environments. Nevertheless, they encourage further testing to determine whether or not the identification of plant groups, plants or plant parts from highly humified peat is possible by applying fen species-specific biomarker proxies.

Effects of CO2 enrichment on photosynthesis, growth, and nitrogen metabolism of the seagrass Zostera noltii

Seagrass ecosystems are expected to benefit from the global increase in CO2 in the ocean because the photosynthetic rate of these plants may be Ci-limited at the current CO2 level. As well, it is expected that lower external pH will facilitate the nitrate uptake of seagrasses if nitrate is cotransported with H+ across the membrane as in terrestrial plants. Here, we investigate the effects of CO2 enrichment on both carbon and nitrogen metabolism of the seagrass Zostera noltii in a mesocosm experiment where plants were exposed for 5 months to two experimental CO2 concentrations (360 and 700 ppm). Both the maximum photosynthetic rate (Pm) and photosynthetic efficiency (a) were higher (1.3- and 4.1-fold, respectively) in plants exposed to CO2-enriched conditions. On the other hand, no significant effects of CO2 enrichment on leaf growth rates were observed, probably due to nitrogen limitation as revealed by the low nitrogen content of leaves. The leaf ammonium uptake rate and glutamine synthetase activity were not significantly affected by increased CO2 concentrations. On the other hand, the leaf nitrate uptake rate of plants exposed to CO2-enriched conditions was fourfold lower than the uptake of plants exposed to current CO2 level, suggesting that in the seagrass Z. noltii nitrate is not cotransported with H+ as in terrestrial plants. In contrast, the activity of nitrate reductase was threefold higher in plant leaves grown at high-CO2 concentrations. Our results suggest that the global effects of CO2 on seagrass production may be spatially heterogeneous and depend on the specific nitrogen availability of each system. Under a CO2 increase scenario, the natural levels of nutrients will probably become limiting for Z. noltii. This potential limitation becomes more relevant because the expected positive effect of CO2 increase on nitrate uptake rate was not confirmed.

Seawater carbonate chemistry, biomass and metabolic rates (leucine incorporation, CO2 fixation and respiration) of Rhodobacteraceae (strain MED165) and Flavobacteriaceae (strain MED217) in a laboratory experiment

Experimental results related to the effects of ocean acidification on planktonic marine microbes are still rather inconsistent and occasionally contradictory. Moreover, laboratory or field experiments that address the effects of changes in CO2 concentrations on heterotrophic microbes are very scarce, despite the major role of these organisms in the marine carbon cycle. We tested the direct effect of an elevated CO2 concentration (1000 ppmv) on the biomass and metabolic rates (leucine incorporation, CO2 fixation and respiration) of 2 isolates belonging to 2 relevant marine bacterial families, Rhodobacteraceae (strain MED165) and Flavobacteriaceae (strain MED217). Our results demonstrate that, contrary to some expectations, high pCO2 did not negatively affect bacterial growth but increased growth efficiency in the case of MED217. The elevated partial pressure of CO2 (pCO2) caused, in both cases, higher rates of CO2 fixation in the dissolved fraction and, in the case of MED217, lower respiration rates. Both responses would tend to increase the pH of seawater acting as a negative feedback between elevated atmospheric CO2 concentrations and ocean acidification.

Seawater carbonate chemistry, calcification rate, oxygen production, maximum quantum yield, symbiont density, chlorophyll concentration and crystal width of Halimeda macroloba, Halimeda cylindracea and Marginopora vertebralis during experiments, 2011

The effects of elevated CO2 and temperature on photosynthesis and calcification in the calcifying algae Halimeda macroloba and Halimeda cylindracea and the symbiont-bearing benthic foraminifera Marginopora vertebralis were investigated through exposure to a combination of four temperatures (28°C, 30°C, 32°C, and 34°C) and four CO2 levels (39, 61, 101, and 203 Pa; pH 8.1, 7.9, 7.7, and 7.4, respectively). Elevated CO2 caused a profound decline in photosynthetic efficiency (FV : FM), calcification, and growth in all species. After five weeks at 34°C under all CO2 levels, all species died. Chlorophyll (Chl) a and b concentration in Halimeda spp. significantly decreased in 203 Pa, 32°C and 34°C treatments, but Chl a and Chl c2 concentration in M. vertebralis was not affected by temperature alone, with significant declines in the 61, 101, and 203 Pa treatments at 28°C. Significant decreases in FV : FM in all species were found after 5 weeks of exposure to elevated CO2 (203 Pa in all temperature treatments) and temperature (32°C and 34°C in all pH treatments). The rate of oxygen production declined at 61, 101, and 203 Pa in all temperature treatments for all species. The elevated CO2 and temperature treatments greatly reduced calcification (growth and crystal size) in M. vertebralis and, to a lesser extent, in Halimeda spp. These findings indicate that 32°C and 101 Pa CO2, are the upper limits for survival of these species on Heron Island reef, and we conclude that these species will be highly vulnerable to the predicted future climate change scenarios of elevated temperature and ocean acidification.

Combined effects of CO2 and light on large and small isolates of the unicellular N2-fixing cyanobacterium Crocosphaera watsonii from the western tropical Atlantic Ocean

We examined the combined effects of light and pCO2 on growth, CO2-fixation and N2-fixation rates by strains of the unicellular marine N2-fixing cyanobacterium Crocosphaera watsonii with small (WH0401) and large (WH0402) cells that were isolated from the western tropical Atlantic Ocean. In low-pCO2-acclimated cultures (190 ppm) of WH0401, growth, CO2-fixation and N2-fixation rates were significantly lower than those in cultures acclimated to higher (present-day 385 ppm, or future 750 ppm) pCO2 treatments. Growth rates were not significantly different, however, in low-pCO2-acclimated cultures of WH0402 in comparison with higher pCO2 treatments. Unlike previous reports for C. watsonii (strain WH8501), N2-fixation rates did not increase further in cultures of WH0401 or WH0402 when acclimated to 750 ppm relative to those maintained at present-day pCO2. Both light and pCO2 had a significant negative effect on gross : net N2-fixation rates in WH0402 and trends were similar in WH0401, implying that retention of fixed N was enhanced under elevated light and pCO2. These data, along with previously reported results, suggest that C. watsonii may have wide-ranging, strain-specific responses to changing light and pCO2, emphasizing the need for examining the effects of global change on a range of isolates within this biogeochemically important genus. In general, however, our data suggest that cellular N retention and CO2-fixation rates of C. watsonii may be positively affected by elevated light and pCO2 within the next 100 years, potentially increasing trophic transfer efficiency of C and N and thereby facilitating uptake of atmospheric carbon by the marine biota.

Effects of ultraviolet radiation and CO2 increase on winter phytoplankton assemblages in a temperate coastal lagoon

Increases in ultraviolet radiation (UVR) and CO2 affect phytoplankton growth and mortality in a variety of different ways. However, in situ responses of natural phytoplankton communities to climate change, as well as its effects on phytoplankton annual cycles, are still largely unknown. Although temperature and UVR have been increasing in temperate latitudes during winter, this season is still particularly neglected in climate change studies, being considered a non-active season regarding phytoplankton growth and production. Additionally, coastal lagoons are highly productive ecosystems and very vulnerable to climate change. This study aims, therefore, to evaluate the short-term effects of increased UVR and CO2 on the composition and growth of winter phytoplankton assemblages in a temperate coastal lagoon. During winter 2012, microcosm experiments were used to evaluate the isolated and combined effects of UVR and CO2, under ambient and high CO2 treatments, exposed to ambient UV levels and photosynthetically active radiation (PAR), or to PAR only. Phytoplankton composition, abundance, biomass and photosynthetic parameters were evaluated during the experiments. Significant changes were observed in the growth of specific phytoplankton groups, leading to changes in community composition. The cyanobacterium Synechococcus was dominant at the beginning of the experiment, but it was negatively affected by UVR and CO2. Diatoms clearly benefited from high CO2 and UVR, particularly Thalassiosira. Despite the changes observed in specific phytoplankton groups, growth and production of the whole phytoplankton community did not show significant responses to UVR and/or CO2.

Seawater carbonate chemistry and Amphiprion melanopus activity during experiments, 2011

Two of the major threats to coral reefs are increasing sea surface temperature and ocean acidification, both of which result from rising concentrations of atmospheric carbon dioxide (CO2). Recent evidence suggests that both increased water temperature and elevated levels of dissolved CO2 can change the behaviors of fishes in ways that reduce individual fitness, however the interacting effects of these variables are unknown. We used a fully factorial experiment to test the independent and interactive effects of temperature (3 levels: 28.5, 30, and 31.5 °C) and pCO2 (3 levels: averaging 420, 530, and 960 µatm) on food consumption and activity level of juvenile anemonefish Amphiprion melanopus (Bleeker 1852). Experimental levels were consistent with current-day ocean conditions and predictions for mid-century and late-century based on atmospheric CO2 projections. Sibling fish were reared for 21 days from the end of their larval phase in each of the nine treatments, at which time behavioral observations were conducted. Food consumption and foraging activity decreased at the highest temperature. In isolation, CO2 level did not significantly affect behavior; however, there was an interaction with temperature. While rearing at high temperature (31.5 °C) and control (420 µatm) or moderate (530 µatm) CO2 resulted in a reduction of food consumption and foraging activity, rearing at high temperature and high CO2 (960 µatm) resulted in an elevation in these behaviors. Maintaining food consumption and foraging activity in high temperature and CO2 conditions may reduce energy efficiency if the thermal optimum for food assimilation and growth has been exceeded. Maintaining foraging effort might increase predation vulnerability. These results suggest that changes in foraging behaviors caused by the interactive effects of increased SST and CO2 could have significant effects on the growth and survival of juvenile reef fishes by late century.

Physiological response to elevated temperature and pCO2 varies across four Pacific coral species: Understanding the unique host + symbiont response

The physiological response to individual and combined stressors of elevated temperature and pCO2 were measured over a 24-day period in four Pacific corals and their respective symbionts (Acropora millepora/Symbiodinium C21a, Pocillopora damicornis/Symbiodinium C1c-d-t, Montipora monasteriata/Symbiodinium C15, and Turbinaria reniformis/Symbiodinium trenchii). Multivariate analyses indicated that elevated temperature played a greater role in altering physiological response, with the greatest degree of change occurring within M. monasteriata and T. reniformis. Algal cellular volume, protein, and lipid content all increased for M. monasteriata. Likewise, S. trenchii volume and protein content in T. reniformis also increased with temperature. Despite decreases in maximal photochemical efficiency, few changes in biochemical composition (i.e. lipids, proteins, and carbohydrates) or cellular volume occurred at high temperature in the two thermally sensitive symbionts C21a and C1c-d-t. Intracellular carbonic anhydrase transcript abundance increased with temperature in A. millepora but not in P. damicornis, possibly reflecting differences in host mitigated carbon supply during thermal stress. Importantly, our results show that the host and symbiont response to climate change differs considerably across species and that greater physiological plasticity in response to elevated temperature may be an important strategy distinguishing thermally tolerant vs. thermally sensitive species.

Effects of ocean acidification on Posidonia oceanica epiphytic community and shoot productivity

1. Biological interactions can alter predictions that are based on single-species physiological response. It is known that leaf segments of the seagrass Posidonia oceanica will increase photosynthesis with lowered pH, but it is not clear whether the outcome will be altered when the whole plant and its epiphyte community, with different respiratory and photosynthetic demands, are included. In addition, the effects on the Posidonia epiphyte community have rarely been tested under controlled conditions, at near-future pH levels. 2. In order to better evaluate the effects of pH levels as projected for the upcoming decades on seagrass meadows, shoots of P. oceanica with their associated epiphytes were exposed in the laboratory to three pH levels (ambient: 8.1, 7.7 and 7.3, on the total scale) for 4 weeks. Net productivity, respiration, net calcification and leaf fluorescence were measured on several occasions. At the end of the study, epiphyte community abundance and composition, calcareous mass and crustose coralline algae growth were determined. Finally, photosynthesis vs. irradiance curves (PE) was produced from segments of secondary leaves cleaned of epiphytes and pigments extracted. 3. Posidonia leaf fluorescence and chlorophyll concentrations did not differ between pH treatments. Net productivity of entire shoots and epiphyte-free secondary leaves increased significantly at the lowest pH level yet limited or no stimulation in productivity was observed at the intermediate pH treatment. Under both pH treatments, significant decreases in epiphytic cover were observed, mostly due to the reduction of crustose coralline algae. The loss of the dominant epiphyte producer yet similar photosynthetic response for epiphyte-free secondary leaves and shoots suggests a minimal contribution of epiphytes to shoot productivity under experimental conditions. 4. Synthesis. Observed responses indicate that under future ocean acidification conditions foreseen in the next century an increase in Posidonia productivity is not likely despite the partial loss of epiphytic coralline algae which are competitors for light. A decline in epiphytic cover could, however, reduce the feeding capacity of the meadow for invertebrates. In situ long-term experiments that consider both acidification and warming scenarios are needed to improve ecosystem-level predictions.

Microenvironmental changes support evidence of photosynthesis and calcification inhibition in Halimeda under ocean acidification and warming

The effects of elevated CO2 and temperature on photosynthesis and calcification of two important calcifying reef algae (Halimeda macroloba and Halimeda cylindracea) were investigated with O2 microsensors and chlorophyll a fluorometry through a combination of two pCO2 (400 and 1,200 µatm) and two temperature treatments (28 and 32 °C) equivalent to the present and predicted conditions during the 2100 austral summer. Combined exposure to pCO2 and elevated temperature impaired calcification and photosynthesis in the two Halimeda species due to changes in the microenvironment around the algal segments and a reduction in physiological performance. There were no significant changes in controls over the 5-week experiment, but there was a 50-70 % decrease in photochemical efficiency (maximum quantum yield), a 70-80 % decrease in O2 production and a threefold reduction in calcification rate in the elevated CO2 and high temperature treatment. Calcification in these species is closely coupled with photosynthesis, such that a decrease in photosynthetic efficiency leads to a decrease in calcification. Although pH seems to be the main factor affecting Halimeda species, heat stress also has an impact on their photosystem II photochemical efficiency. There was a strong combined effect of elevated CO2 and temperature in both species, where exposure to elevated CO2 or temperature alone decreased photosynthesis and calcification, but exposure to both elevated CO2 and temperature caused a greater decline in photosynthesis and calcification than in each stress individually. Our study shows that ocean acidification and ocean warming are drivers of calcification and photosynthesis inhibition in Halimeda. Predicted climate change scenarios for 2100 would therefore severely affect the fitness of Halimeda, which can result in a strongly reduced production of carbonate sediments on coral reefs under such changed climate conditions.

Seawater carbonate chemistry and physiological response of the Mediterranean crustose coralline alga Lithophyllum cabiochae to elevated pCO2 and temperature

The response of respiration, photosynthesis, and calcification to elevated pCO2 and temperature was investigated in isolation and in combination in the Mediterranean crustose coralline alga Lithophyllum cabiochae. Algae were maintained in aquaria during 1 year at near-ambient conditions of irradiance, at ambient or elevated temperature (+3 °C), and at ambient (ca. 400 µatm) or elevated pCO2 (ca. 700 µatm). Respiration, photosynthesis, and net calcification showed a strong seasonal pattern following the seasonal variations of temperature and irradiance, with higher rates in summer than in winter. Respiration was unaffected by pCO2 but showed a general trend of increase at elevated temperature at all seasons, except in summer under elevated pCO2. Conversely, photosynthesis was strongly affected by pCO2 with a decline under elevated pCO2 in summer, autumn, and winter. In particular, photosynthetic efficiency was reduced under elevated pCO2. Net calcification showed different responses depending on the season. In summer, net calcification increased with rising temperature under ambient pCO2 but decreased with rising temperature under elevated pCO2. Surprisingly, the highest rates in summer were found under elevated pCO2 and ambient temperature. In autumn, winter, and spring, net calcification exhibited a positive or no response at elevated temperature but was unaffected by pCO2. The rate of calcification of L. cabiochae was thus maintained or even enhanced under increased pCO2. However, there is likely a trade-off with other physiological processes. For example, photosynthesis declines in response to increased pCO2 under ambient irradiance. The present study reports only on the physiological response of healthy specimens to ocean warming and acidification, however, these environmental changes may affect the vulnerability of coralline algae to other stresses such as pathogens and necroses that can cause major dissolution, which would have critical consequence for the sustainability of coralligenous habitats and the budgets of carbon and calcium carbonate in coastal Mediterranean ecosystems.

Contrasting effects of ocean acidification on tropical fleshy and calcareous algae

Despite the heightened awareness of ocean acidification (OA) effects on marine organisms, few studies empirically juxtapose biological responses to CO2 manipulations across functionally distinct primary producers, particularly benthic algae. Algal responses to OA may vary because increasing CO2 has the potential to fertilize photosynthesis but impair biomineralization. Using a series of repeated experiments on Palmyra Atoll, simulated OA effects were tested across a suite of ecologically important coral reef algae, including five fleshy and six calcareous species. Growth, calcification and photophysiology were measured for each species independently and metrics were combined from each experiment using a meta-analysis to examine overall trends across functional groups categorized as fleshy, upright calcareous, and crustose coralline algae (CCA). The magnitude of the effect of OA on algal growth response varied by species, but the direction was consistent within functional groups. Exposure to OA conditions generally enhanced growth in fleshy macroalgae, reduced net calcification in upright calcareous algae, and caused net dissolution in CCA. Additionally, three of the five fleshy seaweeds tested became reproductive upon exposure to OA conditions. There was no consistent effect of OA on algal photophysiology. Our study provides experimental evidence to support the hypothesis that OA will reduce the ability of calcareous algae to biomineralize. Further, we show that CO2 enrichment either will stimulate population or somatic growth in some species of fleshy macroalgae. Thus, our results suggest that projected OA conditions may favor non-calcifying algae and influence the relative dominance of fleshy macroalgae on reefs, perpetuating or exacerbating existing shifts in reef community structure.

Strict and non-strict independent and-parallelism in logic programs: Correctness, efficiency, and compile-time conditions.

This paper presents some fundamental properties of independent and-parallelism and extends its applicability by enlarging the class of goals eligible for parallel execution. A simple model of (independent) and-parallel execution is proposed and issues of correctness and efficiency discussed in the light of this model. Two conditions, "strict" and "non-strict" independence, are defined and then proved sufficient to ensure correctness and efñciency of parallel execution: if goals which meet these conditions are executed in parallel the solutions obtained are the same as those produced by standard sequential execution. Also, in absence of failure, the parallel proof procedure does not genérate any additional work (with respect to standard SLD-resolution) while the actual execution time is reduced. Finally, in case of failure of any of the goals no slow down will occur. For strict independence the results are shown to hold independently of whether the parallel goals execute in the same environment or in sepárate environments. In addition, a formal basis is given for the automatic compile-time generation of independent and-parallelism: compile-time conditions to efficiently check goal independence at run-time are proposed and proved sufficient. Also, rules are given for constructing simpler conditions if information regarding the binding context of the goals to be executed in parallel is available to the compiler.