Adult acclimation to combined temperature and pH stressors significantly enhances reproductive outcomes compared to short-term exposures

This study examined the effects of long-term culture under altered conditions on the Antarctic sea urchin, Sterechinus neumayeri. Sterechinus neumayeri was cultured under the combined environmental stressors of lowered pH (-0.3 and -0.5 pH units) and increased temperature (+2 °C) for 2 years. This time-scale covered two full reproductive cycles in this species and analyses included studies on both adult metabolism and larval development. Adults took at least 6-8 months to acclimate to the altered conditions, but beyond this, there was no detectable effect of temperature or pH. Animals were spawned after 6 and 17 months exposure to altered conditions, with markedly different outcomes. At 6 months, the percentage hatching and larval survival rates were greatest in the animals kept at 0 °C under current pH conditions, whilst those under lowered pH and +2 °C performed significantly less well. After 17 months, performance was not significantly different across treatments, including controls. However, under the altered conditions urchins produced larger eggs compared with control animals. These data show that under long-term culture adult S. neumayeri appear to acclimate their metabolic and reproductive physiology to the combined stressors of altered pH and increased temperature, with relatively little measureable effect. They also emphasize the importance of long-term studies in evaluating effects of altered pH, particularly in slow developing marine species with long gonad maturation times, as the effects of altered conditions cannot be accurately evaluated unless gonads have fully matured under the new conditions.

Seawater carbonate chemistry and calcification rate of cold-water coral Lophelia pertusa during experiments, 2011

Ocean acidity has increased by 30% since preindustrial times due to the uptake of anthropogenic CO2 and is projected to rise by another 120% before 2100 if CO2 emissions continue at current rates. Ocean acidification is expected to have wide-ranging impacts on marine life, including reduced growth and net erosion of coral reefs. Our present understanding of the impacts of ocean acidification on marine life, however, relies heavily on results from short-term CO2 perturbation studies. Here we present results from the first long-term CO2 perturbation study on the dominant reef-building cold-water coral Lophelia pertusa and relate them to results from a short-term study to compare the effect of exposure time on the coral's responses. Short-term (one week) high CO2 exposure resulted in a decline of calcification by 26-29% for a pH decrease of 0.1 units and net dissolution of calcium carbonate. In contrast, L. pertusa was capable to acclimate to acidified conditions in long-term (six months) incubations, leading to even slightly enhanced rates of calcification. Net growth is sustained even in waters sub-saturated with respect to aragonite. Acclimation to seawater acidification did not cause a measurable increase in metabolic rates. This is the first evidence of successful acclimation in a coral species to ocean acidification, emphasizing the general need for long-term incubations in ocean acidification research. To conclude on the sensitivity of cold-water coral reefs to future ocean acidification further ecophysiological studies are necessary which should also encompass the role of food availability and rising temperatures.

Strong shift from HCO3- to CO2 uptake in Emiliania huxleyi with acidification: new approach unravels acclimation versus short-term pH effects

Effects of ocean acidification on Emiliania huxleyi strain RCC 1216 (calcifying, diploid life-cycle stage) and RCC 1217 (non-calcifying, haploid life-cycle stage) were investigated by measuring growth, elemental composition, and production rates under different pCO2 levels (380 and 950 µatm). In these differently acclimated cells, the photosynthetic carbon source was assessed by a (14)C disequilibrium assay, conducted over a range of ecologically relevant pH values (7.9-8.7). In agreement with previous studies, we observed decreased calcification and stimulated biomass production in diploid cells under high pCO2, but no CO2-dependent changes in biomass production for haploid cells. In both life-cycle stages, the relative contributions of CO2 and HCO3 (-) uptake depended strongly on the assay pH. At pH values =< 8.1, cells preferentially used CO2 (>= 90 % CO2), whereas at pH values >= 8.3, cells progressively increased the fraction of HCO3 (-) uptake (~45 % CO2 at pH 8.7 in diploid cells; ~55 % CO2 at pH 8.5 in haploid cells). In contrast to the short-term effect of the assay pH, the pCO2 acclimation history had no significant effect on the carbon uptake behavior. A numerical sensitivity study confirmed that the pH-modification in the (14)C disequilibrium method yields reliable results, provided that model parameters (e.g., pH, temperature) are kept within typical measurement uncertainties. Our results demonstrate a high plasticity of E. huxleyi to rapidly adjust carbon acquisition to the external carbon supply and/or pH, and provide an explanation for the paradoxical observation of high CO2 sensitivity despite the apparently high HCO3 (-) usage seen in previous studies.

Seawater carbonate chemistry, growth rate and light sensitivity of marine diatom Chaetoceros muelleri (strain CSIRO CS-176) during experiments, 2011

This study investigated the impact of photon flux and elevated CO2 concentrations on growth and photosynthetic electron transport on the marine diatom Chaetoceros muelleri and looked for evidence for the presence of a CO2-concentrating mechanism (CCM). pH drift experiments clearly showed that C. muelleri has the capacity to use bicarbonate to acquire inorganic carbon through one or multiple CCMs. The final pH achieved in unbuffered cultures was not changed by light intensity, even under very low photon flux, implying a low energy demand of bicarbonate use via a CCM. In short-term pH drift experiments, only treatment with the carbonic anhydrase inhibitor ethoxyzolamide (EZ) slowed down the rise in pH considerably. EZ was also the only inhibitor that altered the final pH attained, although marginally. In growth experiments, CO2 availability was manipulated by changing the pH in closed flasks at a fixed dissolved inorganic carbon (DIC) concentration. Low-light-treated samples showed lower growth rates in elevated CO2conditions. No CO2 effect was recorded under high light exposure. The maximal photosynthetic capacity, however, increased with CO2 concentration in saturating, but not in subsaturating, light intensities. Growth and photosynthetic capacity therefore responded in opposite ways to increasing CO2 availability. The capacity to photoacclimate to high and low photon flux appeared not to be affected by CO2treatments. However, photoacclimation was restricted to growth photon fluxes between 30 and 300 µmol photons m-2 s-1. The light saturation points for photosynthetic electron transport and for growth coincided at 100 µmol photons m-2 s-1. Below 100 µmol photons m-2 s-1 the light saturation point for photosynthesis was higher than the growth photon flux (i.e. photosynthesis was not light saturated under growth conditions), whereas at higher growth photon flux, photosynthesis was saturated below growth light levels.

Fast coral reef calcifiers are more sensitive to ocean acidification in short-term laboratory incubations

To identify the properties of taxa sensitive and resistant to ocean acidification (OA), we tested the hypothesis that coral reef calcifiers differ in their sensitivity to OA as predictable outcomes of functional group alliances determined by conspicuous traits. We contrasted functional groups of eight corals and eight calcifying algae defined by morphology in corals and algae, skeletal structure in corals, spatial location of calcification in algae, and growth rate in corals and algae. The responses of calcification to OA were unrelated to morphology and skeletal structure in corals; they were, however, affected by growth rate in corals and algae (fast calcifiers were more sensitive than slow calcifiers), and by the site of calcification and morphology in algae. Species assemblages characterized by fast growth, and for algae, also cell-wall calcification, are likely to be ecological losers in the future ocean. This shift in relative success will affect the relative and absolute species abundances as well as the goods and services provided by coral reefs.

Short- versus long-term responses to changing CO2 in a coastal dinoflagellate bloom

Increasing pCO2 (partial pressure of CO2 ) in an "acidified" ocean will affect phytoplankton community structure, but manipulation experiments with assemblages briefly acclimated to simulated future conditions may not accurately predict the long-term evolutionary shifts that could affect inter-specific competitive success. We assessed community structure changes in a natural mixed dinoflagellate bloom incubated at three pCO2 levels (230, 433, and 765 ppm) in a short-term experiment (2 weeks). The four dominant species were then isolated from each treatment into clonal cultures, and maintained at all three pCO2 levels for approximately 1 year. Periodically (4, 8, and 12 months), these pCO2 -conditioned clones were recombined into artificial communities, and allowed to compete at their conditioning pCO2 level or at higher and lower levels. The dominant species in these artificial communities of CO2 -conditioned clones differed from those in the original short-term experiment, but individual species relative abundance trends across pCO2 treatments were often similar. Specific growth rates showed no strong evidence for fitness increases attributable to conditioning pCO2 level. Although pCO2 significantly structured our experimental communities, conditioning time and biotic interactions like mixotrophy also had major roles in determining competitive outcomes. New methods of carrying out extended mixed species experiments are needed to accurately predict future long-term phytoplankton community responses to changing pCO2 .

Seawater carbonate chemistry, survival, metamorphosis and respiration rate of coral Acropora digitifera during experiments, 2011

Ocean acidification may negatively impact the early life stages of some marine invertebrates including corals. Although reduced growth of juvenile corals in acidified seawater has been reported, coral larvae have been reported to demonstrate some level of tolerance to reduced pH. We hypothesize that the observed tolerance of coral larvae to low pH may be partly explained by reduced metabolic rates in acidified seawater because both calcifying and non-calcifying marine invertebrates could show metabolic depression under reduced pH in order to enhance their survival. In this study, after 3-d and 7-d exposure to three different pH levels (8.0, 7.6, and 7.3), we found that the oxygen consumption of Acropora digitifera larvae tended to be suppressed with reduced pH, although a statistically significant difference was not observed between pH conditions. Larval metamorphosis was also observed, confirming that successful recruitment is impaired when metamorphosis is disrupted, despite larval survival. Results also showed that the metamorphosis rate significantly decreased under acidified seawater conditions after both short (2 h) and long (7 d) term exposure. These results imply that acidified seawater impacts larval physiology, suggesting that suppressed metabolism and metamorphosis may alter the dispersal potential of larvae and subsequently reduce the resilience of coral communities in the near future as the ocean pH decreases.

Nannofossil and planktic foraminiferal events of ODP Site 166-1006 sediments

Detailed biostratigraphy in Site 1006 based on planktonic foraminifers and nannofossils shows large-scale sedimentation rate variability in the Florida Strait west of the Great Bahama Bank. A 'floating' cyclostratigraphy based mainly on resistivity logs and magnetic susceptibility data has been fixed to the biostratigraphy in the absence of magnetostratigraphy. The strongest orbital cycle present is the precessional beat, which is present in the borehole logs throughout the record. Counting the cycles resulted in an accurate time scale and thus a sedimentation rate time series. Spectral analysis of the sedimentation rate time series shows that the short-term cycle of eccentricity (~125 k.y.) and the long term cycle of eccentricity (~400 k.y.) are pervasive throughout the Miocene record, together with the long-term ~2-m.y. eccentricity cycle. The Great Bahama Bank produced pulses of shallow carbonate input once every precessional (sea level) cycle during the Miocene and perhaps two pulses per cycle in the early Pliocene. The amount of sediment exported in these pulses appears to be controlled by eccentricity modulation of the precessional amplitude and therefore the amplitude of the sea-level rise. Finally, an increase in sedimentation rate just after the Miocene/Pliocene boundary is attributed to a change in the location and strength of sediment drift currents in the Florida Strait due to reorganization of the currents following the closure of the Panama Isthmus.

A 18-years long (1993-2011) snow and meteorological dataset from a mid-altitude mountain site (Col de Porte, France, 1325 altitude)

A quality-controlled snow and meteorological dataset spanning the period 1 August 1993-31 July 2011 is presented, originating from the experimental station Col de Porte (1325 m altitude, Chartreuse range, France). Emphasis is placed on meteorological data relevant to the observation and modelling of the seasonal snowpack. In-situ driving data, at the hourly resolution, consist of measurements of air temperature, relative humidity, windspeed, incoming short-wave and long-wave radiation, precipitation rate partitioned between snow- and rainfall, with a focus on the snow-dominated season. Meteorological data for the three summer months (generally from 10 June to 20 September), when the continuity of the field record is not warranted, are taken from a local meteorological reanalysis (SAFRAN), in order to provide a continuous and consistent gap-free record. Data relevant to snowpack properties are provided at the daily (snow depth, snow water equivalent, runoff and albedo) and hourly (snow depth, albedo, runoff, surface temperature, soil temperature) time resolution. Internal snowpack information is provided from weekly manual snowpit observations (mostly consisting in penetration resistance, snow type, snow temperature and density profiles) and from a hourly record of temperature and height of vertically free ''settling'' disks. This dataset has been partially used in the past to assist in developing snowpack models and is presented here comprehensively for the purpose of multi-year model performance assessment. The data is placed on the PANGAEA repository (doi:10.1594/PANGAEA.774249) as well as on the public ftp server ftp://ftp-cnrm.meteo.fr/pub-cencdp/.

Copepod vital rates under CO2-induced acidification: a calanoid species and a cyclopoid species under short-term exposures

Although copepods have been considered tolerant against the direct influence of the ocean acidification (OA) projected for the end of the century, some recent studies have challenged this view. Here, we have examined the direct impact of short-term exposure to a pCO2/pH level relevant for the year 2100 (pHNBS, control: 8.18, low pH: 7.78), on the physiological performance of two representative marine copepods: the calanoid Acartia grani and the cyclopoid Oithona davisae. Adults of both species, from laboratory cultures, were preconditioned for four consecutive days in algal suspensions (Akashiwo sanguinea) prepared with filtered sea water pre-adjusted to the targeted pH values via CO2 bubbling. We measured the feeding and respiratory activity and reproductive output of those pre-conditioned females. The largely unaffected fatty acid composition of the prey offered between OA treatments and controls supports the absence in the study of indirect OA effects (i.e. changes of food nutritional quality). Our results show no direct effect of acidification on the vital rates examined in either copepod species. Our findings are compared with results from previous short- and long-term manipulative experiments on other copepod species.

Seawater carbonate chemistry, survival rate, shell size, calcification rate of the planktonic foraminifer Neogloboquadrina pachyderma (sinistral) in a laboratory experiment

The present study investigated the effects of ocean acidification and temperature increase on Neogloboquadrina pachyderma (sinistral), the dominant planktonic foraminifer in the Arctic Ocean. Due to the naturally low concentration of [CO3] 2- in the Arctic, this foraminifer could be particularly sensitive to the forecast changes in seawater carbonate chemistry. To assess potential responses to ocean acidification and climate change, perturbation experiments were performed on juvenile and adult specimens by manipulating seawater to mimic the present-day carbon dioxide level and a future ocean acidification scenario (end of the century) under controlled (in situ) and elevated temperatures (1 and 4 °C, respectively). Foraminifera mortality was unaffected under all the different experiment treatments. Under low pH, N. pachyderma (s) shell net calcification rates decreased. This decrease was higher (30 %) in the juvenile specimens than decrease observed in the adults (21 %) ones. However, decrease in net calcification was moderated when both, pH decreased and temperature increased simultaneously. When only temperature increased, a net calcification rate for both life stages was not affected. These results show that forecast changes in seawater chemistry would impact calcite production in N. pachyderma (s), possibly leading to a reduction of calcite flux contribution and consequently a decrease in biologic pump efficiency.

Properties of seawater and particulate matter from a FRRF instrument, operating in a flow-through mode on the continuous surface water sampling system during the Tara Oceans expedition 2009-2013

The Tara Oceans Expedition (2009-2013) sampled the world oceans on board a 36 m long schooner, collecting environmental data and organisms from viruses to planktonic metazoans for later analyses using modern sequencing and state-of-the-art imaging technologies. Tara Oceans Data are particularly suited to study the genetic, morphological and functional diversity of plankton. The present data set provides continuous measurements made with a FRRF instrument, operating in a flow-through mode during the 2009-2012 part of the expedition. It operates by exciting chlorophyll fluorescence using a series of short flashes of controlled energy and time intervals (Kolber et al, 1998). The fluorescence transients produced by this excitation signal were analysed in real-time to provide estimates of abundance of photosynthetic pigments, the photosynthetic yields (Fv/Fm), the functional absorption cross section (a proxy for efficiency of photosynthetic energy acquisition), the kinetics of photosynthetic electron transport between Photosystem II and Photosystem I, and the size of the PQ pool. These parameters were measured at excitation wavelength of 445 nm, 470nm, 505 nm, and 535 nm, allowing to assess the presence and the photosynthetic performance of different phytoplankton taxa based on the spectral composition of their light harvesting pigments. The FRRF-derived photosynthetic characteristics were used to calculate the initial slope, the half saturation, and the maximum level of Photosynthesis vs Irradiance relationship. FRRF data were acquired continuously, at 1-minute time intervals.

Photophysiological responses of Southern Ocean phytoplankton to changes in CO2 concentrations: Short-term versus acclimation effects

The present study examines how different pCO2 acclimations affect the CO2- and light-dependence of photophysiological processes and O2 fluxes in four Southern Ocean (SO) key phytoplankton species. We grew Chaetoceros debilis (Cleve), Pseudo-nitzschia subcurvata (Hasle), Fragilariopsis kerguelensis (O'Meara) and Phaeocystis antarctica (Karsten) under low (160 µatm) and high (1000 ?atm) pCO2. The CO2- and light-dependence of fluorescence parameters of photosystem II (PSII) were determined by means of a fluorescence induction relaxation system (FIRe). In all tested species, nonphotochemical quenching (NPQ) is the primary photoprotection strategy in response to short-term exposure to high light or low CO2 concentrations. In C. debilis and P. subcurvata, PSII connectivity (p) and functional absorption cross-sections of PSII in ambient light (sigma PSII') also contributed to photoprotection while changes in re-oxidation times of Qa acceptor (tQa) were more significant in F. kerguelensis. The latter was also the only species being responsive to high acclimation pCO2, as these cells had enhanced relative electron transport rates (rETRs) and sigma PSII' while tQa and p were reduced under short-term exposure to high irradiance. Low CO2-acclimated cells of F. kerguelensis and all pCO2 acclimations of C. debilis and P. subcurvata showed dynamic photoinhibition with increasing irradiance. To test for the role and presence of the Mehler reaction in C. debilis and P. subcurvata, the light-dependence of O2 fluxes was estimated using membrane inlet mass spectrometry (MIMS). Our results show that the Mehler reaction is absent in both species under the tested conditions. We also observed that dark respiration was strongly reduced under high pCO2 in C. debilis while it remained unaltered in P. subcurvata. Our study revealed species-specific differences in the photophysiological responses to pCO2, both on the acclimation as well as the short-term level.