Effects of ocean acidification on the swimming ability, development and biochemical responses of sand smelt larvae

Ocean acidification, recognized as a major threat to marine ecosystems, has developed into one of the fastest growing fields of research in marine sciences. Several studies on fish larval stages point to abnormal behaviours, malformations and increased mortality rates as a result of exposure to increased levels of CO2. However, other studies fail to recognize any consequence, suggesting species-specific sensitivity to increased levels of CO2, highlighting the need of further research. In this study we investigated the effects of exposure to elevated pCO2 on behaviour, development, oxidative stress and energy metabolism of sand smelt larvae, Atherina presbyter. Larvae were caught at Arrábida Marine Park (Portugal) and exposed to different pCO2 levels (control: 600 µatm, pH = 8.03; medium: 1000 µatm, pH = 7.85; high: 1800 µatm, pH = 7.64) up to 15 days, after which critical swimming speed (Ucrit), morphometric traits and biochemical biomarkers were determined. Measured biomarkers were related with: 1) oxidative stress-superoxide dismutase and catalase enzyme activities, levels of lipid peroxidation and DNA damage, and levels of superoxide anion production; 2) energy metabolism - total carbohydrate levels, electron transport system activity, lactate dehydrogenase and isocitrate dehydrogenase enzyme activities. Swimming speed was not affected by treatment, but exposure to increasing levels of pCO2 leads to higher energetic costs and morphometric changes, with larger larvae in high pCO2 treatment and smaller larvae in medium pCO2 treatment. The efficient antioxidant response capacity and increase in energetic metabolism only registered at the medium pCO2 treatment may indicate that at higher pCO2 levels the capacity of larvae to restore their internal balance can be impaired. Our findings illustrate the need of using multiple approaches to explore the consequences of future pCO2 levels on organisms.

Seawater carbonate chemistry and biological processes during experiments with early life stages of the blue mussel Mytilus edulis, 2010

Several experiments have shown a decrease of growth and calcification of organisms at decreased pH levels. There is a growing interest to focus on early life stages that are believed to be more sensitive to environmental disturbances such as hypercapnia. Here, we present experimental data, acquired in a commercial hatchery, demonstrating that the growth of planktonic mussel (Mytilus edulis) larvae is significantly affected by a decrease of pH to a level expected for the end of the century. Even though there was no significant effect of a 0.25-0.34 pH unit decrease on hatching and mortality rates during the first 2 days of development nor during the following 13-day period prior to settlement, final shells were respectively 4.5±1.3 and 6.0±2.3% smaller at pHNBS~7.8 (pCO2~1100-1200 µatm) than at a control pHNBS of ~8.1 (pCO2~460-640 µatm). Moreover, a decrease of 12.0±5.4% of shell thickness was observed after 15d of development. More severe impacts were found with a decrease of ~0.5 pHNBS unit during the first 2 days of development which could be attributed to a decrease of calcification due to a slight undersaturation of seawater with respect to aragonite. Indeed, important effects on both hatching and D-veliger shell growth were found. Hatching rates were 24±4% lower while D-veliger shells were 12.7±0.9% smaller at pHNBS~7.6 (pCO2~1900 µatm) than at a control pHNBS of ~8.1 (pCO2~540 µatm). Although these results show that blue mussel larvae are still able to develop a shell in seawater undersaturated with respect to aragonite, the observed decreases of hatching rates and shell growth could lead to a significant decrease of the settlement success. As the environmental conditions considered in this study do not necessarily reflect the natural conditions experienced by this species at the time of spawning, future studies will need to consider the whole larval cycle (from fertilization to settlement) under environmentally relevant conditions in order to investigate the potential ecological and economical losses of a decrease of this species fitness in the field.

Nighttime dissolution in a temperate coastal ocean ecosystem increases under acidification

Anthropogenic emissions of carbon dioxide (CO2) are causing ocean acidification, lowering seawater aragonite (CaCO3) saturation state (Omega arag), with potentially substantial impacts on marine ecosystems over the 21st Century. Calcifying organisms have exhibited reduced calcification under lower saturation state conditions in aquaria. However, the in situ sensitivity of calcifying ecosystems to future ocean acidification remains unknown. Here we assess the community level sensitivity of calcification to local CO2-induced acidification caused by natural respiration in an unperturbed, biodiverse, temperate intertidal ecosystem. We find that on hourly timescales nighttime community calcification is strongly influenced by Omega arag, with greater net calcium carbonate dissolution under more acidic conditions. Daytime calcification however, is not detectably affected by Omega arag. If the short-term sensitivity of community calcification to Omega arag is representative of the long-term sensitivity to ocean acidification, nighttime dissolution in these intertidal ecosystems could more than double by 2050, with significant ecological and economic consequences.

Seawater carbonate chemistry and processes during experiments with phytoplankton Emiliania huxleyi (strain Bergen 2005), 2010

The response of the coccolithophore Emiliania huxleyi to rising CO2 concentrations is well documented for acclimated cultures where cells are exposed to the CO2 treatments for several generations prior to the experiment. The exact number of generations required for acclimation to CO2-induced changes in seawater carbonate chemistry, however, is unknown. Here we show that Emiliania huxleyi's short-term response (26 h) after cultures (grown at 500 µatm) were abruptly exposed to changed CO2 concentrations (~190, 410, 800 and 1500 ?atm) is similar to that obtained with acclimated cultures under comparable conditions in earlier studies. Most importantly, from the lower CO2 levels (190 and 410 ?atm) to 750 and 1500 µatm calcification decreased and organic carbon fixation increased within the first 8 to 14 h after exposing the cultures to changes in carbonate chemistry. This suggests that Emiliania huxleyi rapidly alters the rates of essential metabolical processes in response to changes in seawater carbonate chemistry, establishing a new physiological "state" (acclimation) within a matter of hours. If this relatively rapid response applies to other phytoplankton species, it may simplify interpretation of studies with natural communities (e.g. mesocosm studies and ship-board incubations), where often it is not feasible to allow for a pre-conditioning phase before starting experimental incubations.

Seawater carbonate chemistry and processes during experiments with Emiliania huxleyi (2005 Bergen) and Coccolithus braarudii (RCC 1200), 2010

The physiological performance of two coccolithophore species,Emiliania huxleyi and Coccolithus braarudii, was investigated during long-term exposure to elevated pCO2 levels. Mono-specific cultures were grown over 152 (E. huxleyi) and 65 (C. braarudii) generations while pCO2 was gradually increased to maximum levels of 1150 ?atm (E. huxleyi) and 930 ?atm (C. braarudii) and kept constant thereafter. Rates of cell growth and cell quotas of particulate organic carbon (POC), particulate inorganic carbon (PIC) and total particulate nitrogen (TPN) were determined repeatedly throughout the incubation period. Increasing pCO2 caused a decrease in cell growth rate of 9% and 29% in E. huxleyi and C. braarudii, respectively. In both species cellular PIC:TPN and PIC:POC ratios decreased in response to rising pCO2, whereas no change was observed in the POC:TPN ratios of E. huxleyi and C. braarudii. These results are consistent with those obtained in shorter-term high CO2exposure experiments following abrupt pertubations of the seawater carbonate system and indicate that for the strains tested here a gradual CO2 increase does not alleviate CO2/pH sensitivity.

Seal census raw data of campaigns EMAGE I to V between 1996 and 2000 with links to datasets

The development of models of marine ecosystems in the Southern Ocean is becoming increasingly important as a means of understanding and managing impacts such as exploitation and climate change. Collating data from disparate sources, and understanding biases or uncertainties inherent in those data, are important first steps for improving ecosystem models. This review focuses on seals that breed in ice habitats of the Southern Ocean (i.e. the crabeater seal, Lobodon carcinophaga; Ross seal, Ommatophoca rossii; leopard seal, Hydrurga leptonyx; and Weddell seal, Leptonychotes weddellii). Data on populations (abundance and trends in abundance), distribution and habitat use (movement, key habitat and environmental features) and foraging (diet) are summarised, and potential biases and uncertainties inherent in those data are identified and discussed. Spatial and temporal gaps in knowledge of the populations, habitats and diet of each species are also identified.

Experiment: Metabolic shifts in the Antarctic fish Notothenia rossii in response to rising temperature and PCO2

Introduction Ongoing ocean warming and acidification increasingly affect marine ecosystems, in particular around the Antarctic Peninsula. Yet little is known about the capability of Antarctic notothenioid fish to cope with rising temperature in acidifying seawater. While the whole animal level is expected to be more sensitive towards hypercapnia and temperature, the basis of thermal tolerance is set at the cellular level, with a putative key role for mitochondria. This study therefore investigates the physiological responses of the Antarctic Notothenia rossii after long-term acclimation to increased temperatures (7°C) and elevated PCO2 (0.2 kPa CO2) at different levels of physiological organisation. Results For an integrated picture, we analysed the acclimation capacities of N. rossii by measuring routine metabolic rate (RMR), mitochondrial capacities (state III respiration) as well as intra- and extracellular acid-base status during acute thermal challenges and after long-term acclimation to changing temperature and hypercapnia. RMR was partially compensated during warm- acclimation (decreased below the rate observed after acute warming), while elevated PCO2 had no effect on cold or warm acclimated RMR. Mitochondrial state III respiration was unaffected by temperature acclimation but depressed in cold and warm hypercapnia-acclimated fish. In both cold- and warm-exposed N. rossii, hypercapnia acclimation resulted in a shift of extracellular pH (pHe) towards more alkaline values. A similar overcompensation was visible in muscle intracellular pH (pHi). pHi in liver displayed a slight acidosis after warm normo- or hypercapnia acclimation, nevertheless, long-term exposure to higher PCO2 was compensated for by intracellular bicarbonate accumulation. Conclusion The partial warm compensation in whole animal metabolic rate indicates beginning limitations in tissue oxygen supply after warm-acclimation of N. rossii. Compensatory mechanisms of the reduced mitochondrial capacities under chronic hypercapnia may include a new metabolic equilibrium to meet the elevated energy demand for acid-base regulation. New set points of acid-base regulation under hypercapnia, visible at the systemic and intracellular level, indicate that N. rossii can at least in part acclimate to ocean warming and acidification. It remains open whether the reduced capacities of mitochondrial energy metabolism are adaptive or would impair population fitness over longer timescales under chronically elevated temperature and PCO2.

Seawater carbonate chemistry and sea urchin Paracentrotus lividus biological processes during experiments, 2011

Ocean acidification is predicted to have significant effects on benthic calcifying invertebrates, in particular on their early developmental stages. Echinoderm larvae could be particularly vulnerable to decreased pH, with major consequences for adult populations. The objective of this study was to understand how ocean acidification would affect the initial life stages of the sea urchin Paracentrotus lividus, a common species that is widely distributed in the Mediterranean Sea and the NE Atlantic. The effects of decreased pH (elevated PCO2) were investigated through physiological and molecular analyses on both embryonic and larval stages. Eggs and larvae were reared in Mediterranean seawater at six pH levels, i.e. pHT 8.1, 7.9, 7.7, 7.5, 7.25 and 7.0. Fertilization success, survival, growth and calcification rates were monitored over a 3 day period. The expression of genes coding for key proteins involved in development and biomineralization was also monitored. Paracentrotus lividus appears to be extremely resistant to low pH, with no effect on fertilization success or larval survival. Larval growth was slowed when exposed to low pH but with no direct impact on relative larval morphology or calcification down to pHT 7.25. Consequently, at a given time, larvae exposed to low pH were present at a normal but delayed larval stage. More surprisingly, candidate genes involved in development and biomineralization were upregulated by factors of up to 26 at low pH. Our results revealed plasticity at the gene expression level that allows a normal, but delayed, development under low pH conditions.

Response of A. aurita ephyrae to three stressors: high-temperature, high-CO2 and low-oxygen

Global change is affecting marine ecosystems through a combination of different stressors such as warming, ocean acidification and oxygen depletion. Very little is known about the interactions among these factors, especially with respect to gelatinous zooplankton. Therefore, in this study we investigated the direct effects of pH, temperature and oxygen availability on the moon jellyfish Aurelia aurita, concentrating on the ephyral life stage. Starved one-day-old ephyrae were exposed to a range of pCO2 (400-4000 ppm) and three different dissolved oxygen levels (from saturated to hypoxic conditions), in two different temperatures (5 and 15 °C) for 7 days. Carbon content and swimming activity were analysed at the end of the incubation period, and mortality noted. General linearized models were fitted through the data, with the best fitting models including two- and three-way interactions between pCO2, temperature and oxygen concentration. The combined effect of the stressors was small but significant, with the clearest negative effect on growth caused by the combination of all three stressors present (high temperature, high CO2, low oxygen). We conclude that A. aurita ephyrae are robust and that they are not likely to suffer from these environmental stressors in a near future.

CFC, and noble gas measurements during POLARSTERN cruise ANT-XXIX/3

During the austral summer expedition PS81, ANT-XXIX/3 with the German research ice breaker Polarstern in 2013, research was carried out to investigate the role of environmental factors on the distribution of benthic communities and marine mammal and krill densities around the northern tip of the Antarctic Peninsula. For these studies collated in this special issue and studies in this area, we present a collection of environmental parameters with probable influence on the marine ecosystems around the Antarctic Peninsula.

Seawater carbonate chemistry and hatch rates of Antarctic krill

Marine ecosystems of the Southern Ocean are particularly vulnerable to ocean acidification. Antarctic krill (Euphausia superba; hereafter krill) is the key pelagic species of the region and its largest fishery resource. There is therefore concern about the combined effects of climate change, ocean acidification and an expanding fishery on krill and ultimately, their dependent predators-whales, seals and penguins. However, little is known about the sensitivity of krill to ocean acidification. Juvenile and adult krill are already exposed to variable seawater carbonate chemistry because they occupy a range of habitats and migrate both vertically and horizontally on a daily and seasonal basis. Moreover, krill eggs sink from the surface to hatch at 700-1,000 m, where the carbon dioxide partial pressure (pCO2) in sea water is already greater than it is in the atmosphere. Krill eggs sink passively and so cannot avoid these conditions. Here we describe the sensitivity of krill egg hatch rates to increased CO2, and present a circumpolar risk map of krill hatching success under projected pCO2 levels. We find that important krill habitats of the Weddell Sea and the Haakon VII Sea to the east are likely to become high-risk areas for krill recruitment within a century. Furthermore, unless CO2 emissions are mitigated, the Southern Ocean krill population could collapse by 2300 with dire consequences for the entire ecosystem.

Table 4. Trace gas concentrations from samples taken at Ginninderra

We assess the performance of an inverse Lagrangian dispersion technique for its suitability to quantify leakages from geological storage of CO2. We find the technique is accurate ((QbLS/Q)=0.99, sigma=0.29) when strict meteorological filtering is applied to ensure that Monin-Obukhov Similarity Theory is valid for the periods analysed and when downwind enrichments in tracer gas concentration are 1% or more above background concentration. Because of their respective baseline atmospheric concentrations, this enrichment criterion is less onerous for CH4 than for CO2. Therefore for geologically sequestered gas reservoirs with a significant CH4 component, monitoring CH4 as a surrogate for CO2 leakage could be as much as 10 times more sensitive than monitoring CO2 alone. Additional recommendations for designing a robust atmospheric monitoring strategy for geosequestration include: continuous concentration data; exact inter-calibration of up- and downwind concentration measurements; use of an array of point concentration sensors to maximise the use of spatial information about the leakage plume; and precise isotope ratio measurement to confirm the source of any concentration elevations detected.

He isotope, U/Th isotope, Calcium carbonate, biogenic opal, rare earth element concentrations, and grain size distribution measured on core-top sediments during SONNE cruise SO202 (INOPEX)

Eolian dust is a significant source of iron and other nutrients that are essential for the health of marine ecosystems and potentially a controlling factor of the high nutrient-low chlorophyll status of the Subarctic North Pacific. We map the spatial distribution of dust input using three different geochemical tracers of eolian dust, 4He, 232Th and rare earth elements, in combination with grain size distribution data, from a set of core-top sediments covering the entire Subarctic North Pacific. Using the suite of geochemical proxies to fingerprint different lithogenic components, we deconvolve eolian dust input from other lithogenic inputs such as volcanic ash, ice-rafted debris, riverine and hemipelagic input. While the open ocean sites far away from the volcanic arcs are dominantly composed of pure eolian dust, lithogenic components other than eolian dust play a more crucial role along the arcs. In sites dominated by dust, eolian dust input appears to be characterized by a nearly uniform grain size mode at ~4 µm. Applying the 230Th-normalization technique, our proxies yield a consistent pattern of uniform dust fluxes of 1-2 g/m**2/yr across the Subarctic North Pacific. Elevated eolian dust fluxes of 2-4 g/m**2/yr characterize the westernmost region off Japan and the southern Kurile Islands south of 45° N and west of 165° E along the main pathway of the westerly winds. The core-top based dust flux reconstruction is consistent with recent estimates based on dissolved thorium isotope concentrations in seawater from the Subarctic North Pacific. The dust flux pattern compares well with state-of-the-art dust model predictions in the western and central Subarctic North Pacific, but we find that dust fluxes are higher than modeled fluxes by 0.5-1 g/m**2/yr in the northwest, northeast and eastern Subarctic North Pacific. Our results provide an important benchmark for biogeochemical models and a robust approach for downcore studies testing dust-induced iron fertilization of past changes in biological productivity in the Subarctic North Pacific.

Seawater carbonate chemistry, and Lithophyllum sp. and Feldmannia spp. coverage and dry weight during experiments, 2009

Climate-driven change represents the cumulative effect of global through local-scale conditions, and understanding their manifestation at local scales can empower local management. Change in the dominance of habitats is often the product of local nutrient pollution that occurs at relatively local scales (i.e. catchment scale), a critical scale of management at which global impacts will manifest. We tested whether forecasted global-scale change [elevated carbon dioxide (CO2) and subsequent ocean acidification] and local stressors (elevated nutrients) can combine to accelerate the expansion of filamentous turfs at the expense of calcifying algae (kelp understorey). Our results not only support this model of future change, but also highlight the synergistic effects of future CO2 and nutrient concentrations on the abundance of turfs. These results suggest that global and local stressors need to be assessed in meaningful combinations so that the anticipated effects of climate change do not create the false impression that, however complex, climate change will produce smaller effects than reality. These findings empower local managers because they show that policies of reducing local stressors (e.g. nutrient pollution) can reduce the effects of global stressors not under their governance (e.g. ocean acidification). The connection between research and government policy provides an example whereby knowledge (and decision making) across local through global scales provides solutions to some of the most vexing challenges for attaining social goals of sustainability, biological conservation and economic development.

Seawater carbonate chemistry, larval settlement and growth rate during experiments with coral Porites astreoides, 2008

In response to the increases in pCO2 projected in the 21st century, adult coral growth and calcification are expected to decrease significantly. However, no published studies have investigated the effect of elevated pCO2 on earlier life history stages of corals. Porites astreoides larvae were collected from reefs in Key Largo, Florida, USA, settled and reared in controlled saturation state seawater. Three saturation states were obtained, using 1 M HCl additions, corresponding to present (380 ppm) and projected pCO2 scenarios for the years 2065 (560 ppm) and 2100 (720 ppm). The effect of saturation state on settlement and post-settlement growth was evaluated. Saturation state had no significant effect on percent settlement; however, skeletal extension rate was positively correlated with saturation state, with ~50% and 78% reductions in growth at the mid and high pCO2 treatments compared to controls, respectively.