Ocean acidification and the loss of phenolic substances in marine plants

Rising atmospheric CO2 often triggers the production of plant phenolics, including many that serve as herbivore deterrents, digestion reducers, antimicrobials, or ultraviolet sunscreens. Such responses are predicted by popular models of plant defense, especially resource availability models which link carbon availability to phenolic biosynthesis. CO2 availability is also increasing in the oceans, where anthropogenic emissions cause ocean acidification, decreasing seawater pH and shifting the carbonate system towards further CO2 enrichment. Such conditions tend to increase seagrass productivity but may also increase rates of grazing on these marine plants. Here we show that high CO2 / low pH conditions of OA decrease, rather than increase, concentrations of phenolic protective substances in seagrasses and eurysaline marine plants. We observed a loss of simple and polymeric phenolics in the seagrass Cymodocea nodosa near a volcanic CO2 vent on the Island of Vulcano, Italy, where pH values decreased from 8.1 to 7.3 and pCO2 concentrations increased ten-fold. We observed similar responses in two estuarine species, Ruppia maritima and Potamogeton perfoliatus, in in situ Free-Ocean-Carbon-Enrichment experiments conducted in tributaries of the Chesapeake Bay, USA. These responses are strikingly different than those exhibited by terrestrial plants. The loss of phenolic substances may explain the higher-than-usual rates of grazing observed near undersea CO2 vents and suggests that ocean acidification may alter coastal carbon fluxes by affecting rates of decomposition, grazing, and disease. Our observations temper recent predictions that seagrasses would necessarily be "winners" in a high CO2 world.

Interactive effects of ocean acidification and nitrogen limitation on the diatom Phaeodactylum tricornutum

Climate change is expected to bring about alterations in the marine physical and chemical environment that will induce changes in the concentration of dissolved CO2 and in nutrient availability. These in turn are expected to affect the physiological performance of phytoplankton. In order to learn how phytoplankton respond to the predicted scenario of increased CO2 and decreased nitrogen in the surface mixed layer, we investigated the diatom Phaeodactylum tricornutum as a model organism. The cells were cultured in both low CO2 (390 µatm) and high CO2 (1000 µatm) conditions at limiting (10 µmol/L) or enriched (110 µmol/L) nitrate concentrations. Our study shows that nitrogen limitation resulted in significant decreases in cell size, pigmentation, growth rate and effective quantum yield of Phaeodactylum tricornutum, but these parameters were not affected by enhanced dissolved CO2 and lowered pH. However, increased CO2 concentration induced higher rETRmax and higher dark respiration rates and decreased the CO2 or dissolved inorganic carbon (DIC) affinity for electron transfer (shown by higher values for K1/2 DIC or K1/2 CO2). Furthermore, the elemental stoichiometry (carbon to nitrogen ratio) was raised under high CO2 conditions in both nitrogen limited and nitrogen replete conditions, with the ratio in the high CO2 and low nitrate grown cells being higher by 45% compared to that in the low CO2 and nitrate replete grown ones. Our results suggest that while nitrogen limitation had a greater effect than ocean acidification, the combined effects of both factors could act synergistically to affect marine diatoms and related biogeochemical cycles in future oceans.

Adaptation of a globally important coccolithophore to ocean warming and acidification

Regulating intracellular pH (pHi) is critical for optimising the metabolic activity of corals, yet mechanisms involved in pH regulation and the buffering capacity within coral cells are not well understood. Our study investigated how the presence of symbiotic dinoflagellates affects the response of pHi to pCO2-driven seawater acidification in cells isolated from Pocillopora damicornis. Using the fluorescent dye BCECF-AM, in conjunction with confocal microscopy, we simultaneously characterised the response of pHi in host coral cells and their dinoflagellate symbionts, in symbiotic and non-symbiotic states under saturating light, with and without the photosynthetic inhibitor DCMU. Each treatment was run under control (pH 7.8) and CO2 acidified seawater conditions (decreasing pH from 7.8 - 6.8). After two hours of CO2 addition, by which time the external pH (pHe) had declined to 6.8, the dinoflagellate symbionts had increased their pHi by 0.5 pH units above control levels. In contrast, in both symbiotic and non-symbiotic host coral cells, 15 min of CO2 addition (0.2 pH unit drop in pHe) led to cytoplasmic acidosis equivalent to 0.4 pH units. Despite further seawater acidification over the duration of the experiment, the pHi of non-symbiotic coral cells did not change, though in host cells containing a symbiont cell the pHi recovered to control levels. This recovery was negated when cells were incubated with DCMU. Our results reveal that photosynthetic activity of the endosymbiont is tightly coupled with the ability of the host cell to recover from cellular acidosis after exposure to high CO2 / low pH.

Impact of ocean acidification and warming on the larval development of the spider crab Hyas araneus from different latitudes (54° vs 79°N), 2010

The combined impacts of future scenarios of ocean acidification and global warming on the larvae of a cold-eurythermal spider crab, Hyas araneus L., were investigated in one of its southernmost populations (living around Helgoland, southern North Sea, 54°N) and one of the northernmost populations (Svalbard, North Atlantic, 79°N). Larvae were exposed at temperatures of 3, 9 and 15°C to present day normocapnia (380 ppm CO2) and to CO2 conditions expected for the near or medium-term future (710 ppm by 2100 and 3000 ppm CO2 by 2300 and beyond). Larval development time and biochemical composition were studied in the larval stages Zoea I, II, and Megalopa. Permanent differences in instar duration between both populations were detected in all stages, likely as a result of evolutionary temperature adaptation. With the exception of Zoea II at 3°C and under all CO2 conditions, development in all instars from Svalbard was delayed compared to those from Helgoland, under all conditions. Most prominently, development was much longer and fewer specimens morphosed to the first crab instar in the Megalopa from Svalbard than from Helgoland. Enhanced CO2 levels (710 and particularly 3000 ppm), caused extended duration of larval development and reduced larval growth (measured as dry mass) and fitness (decreasing C/N ratio, a proxy of the lipid content). Such effects were strongest in the zoeal stages in Svalbard larvae, and during the Megalopa instar in Helgoland larvae.

Combined effects of ocean acidification and light or nitrogen availabilities on 13C fractionation in marine dinoflagellates

Along with increasing oceanic CO2 concentrations, enhanced stratification constrains phytoplankton to shallower upper mixed layers with altered light regimes and nutrient concentrations. Here, we investigate the effects of elevated pCO2 in combination with light or nitrogen-limitation on 13C fractionation (epsilon p) in four dinoflagellate species. We cultured Gonyaulax spinifera and Protoceratium reticulatum in dilute batches under low-light (LL) and high-light (HL) conditions, and grew Alexandrium fundyense and Scrippsiella trochoidea in nitrogen-limited continuous cultures (LN) and nitrogen-replete batches (HN). The observed CO2-dependency of epsilon p remained unaffected by the availability of light for both G. spinifera and P. reticulatum, though at HL epsilon p was consistently lower by about 2.7 per mil over the tested CO2 range for P. reticulatum. This may reflect increased uptake of (13C-enriched) bicarbonate fueled by increased ATP production under HL conditions. The observed CO2-dependency of epsilon p disappeared under LN conditions in both A. fundyense and S. trochoidea. The generally higher epsilon p under LN may be associated with lower organic carbon production rates and/or higher ATP:NADPH ratios. CO2-dependent epsilon p under non-limiting conditions has been observed in several dinoflagellate species, showing potential for a new CO2-proxy. Our results however demonstrate that light- and nitrogen-limitation also affect epsilon p, thereby illustrating the need to carefully consider prevailing environmental conditions.

Interactive effects of ocean acidification and neighboring corals on the growth of Pocillopora verrucosa

The physical and chemical environment around corals, as well as their physiology, can be affected by interactions with neighboring corals. This study employed small colonies (4 cm diameter) of Pocillopora verrucosa and Acropora hyacinthus configured in spatial arrays at 7 cm/s flow speed to test the hypothesis that ocean acidification (OA) alters interactions among them. Interaction effects were quantified for P. verrucosa using three measures of growth: calcification (i.e., weight), horizontal growth, and vertical growth. The study was carried out in May-June 2014 using corals from 10 m depth on the outer reef of Moorea, French Polynesia. Colonies of P. verrucosa were placed next to conspecifics or heterospecifics (A. hyacinthus) in arrangements of two or four colonies (pairs and aggregates) that were incubated at ambient and high pCO2 (1000 µatm) for 28 days. There was an effect of pCO2, and arrangement type on multivariate growth (utilizing the three measures of growth), but no interaction between the main effects. Conversely, arrangement and pCO2 had an interactive effect on calcification, with an overall 23 % depression at high pCO2 versus ambient pCO2 (i.e., pooled among arrangements). Within arrangements, there was a 34-45 % decrease in calcification for solitary and paired conspecifics, but no effect in conspecific aggregates, heterospecific pairs, or heterospecific aggregates. Horizontal growth was negatively affected by pCO2 and arrangement type, while vertical growth was positively affected by arrangement type. Together, our results show that conspecific aggregations can mitigate the negative effects of OA on calcification of colonies within an aggregation.

The synergistic effects of ocean acidification and organic metabolism on calcium carbonate (CaCO3) dissolution in coral reef sediments

Ocean acidification (OA) is expected to reduce the net ecosystem calcification (NEC) rates and overall accretion of coral reef ecosystems. However, despite the fact that sediments are the most abundant form of calcium carbonate (CaCO3) in coral reef ecosystems and their dissolution may be more sensitive to OA than biogenic calcification, the impacts of OA induced sediment dissolution on coral reef NEC rates and CaCO3 accretion are poorly constrained. Carbon dioxide addition and light attenuation experiments were performed at Heron Island, Australia in an attempt to tease apart the influence of OA and organic metabolism (e.g. respiratory CO2 production) on CaCO3 dissolution. Overall, CaCO3 dissolution rates were an order of magnitude more sensitive to elevated CO2 and decreasing seawater aragonite saturation state (Omega Ar; 300-420% increase in dissolution per unit decrease in Omega Ar) than published reductions in biologically mediated calcification due to OA. Light attenuation experiments led to a 70% reduction in net primary production (NPP), which subsequently induced an increase in daytime (115%) and net diel (375%) CaCO3 dissolution rates. High CO2 and low light acted in synergy to drive a 575% increase in net diel dissolution rates. Importantly, disruptions to the balance of photosynthesis and respiration (P/R) had a significant effect on daytime CaCO3 dissolution, while average water column ?Ar was the main driver of nighttime dissolution rates. A simple model of platform-integrated dissolution rates was developed demonstrating that seasonal changes in photosynthetically active radiation (PAR) can have an important effect on platform integrated CaCO3 sediment dissolution rates. The considerable response of CaCO3 sediment dissolution to elevated CO2 means that much of the response of coral reef communities and ecosystems to OA could be due to increases in CaCO3 sediment and framework dissolution, and not decreases in biogenic calcification.

Freak waves and modulational dynamics in plasmas with negative ions

The nonlinear dynamics of modulated electrostatic wavepackets propagating in negativeion plasmas is investigated from first principles. A nonlinear Schrödinger equation is derived by adopting a multiscale technique. The stability of breather- like (bright envelope soliton) structures, considered as a precursor to freak wave (rogue wave) formation, is investigated and then tested via numerical simulations.

Interaction between wind waves and swell in the Persian Gulf

The effect of swell on wind wave growth has been a topic of active research for many years with inconsistent results. The details are often contradictory among investigations. Further more, there remain a variety of competing theories to explain these phenomena. In this research, we consider waves and wind and temperature data in the Persian Gulf (Busher region) in years 1995, 1996 and 1999. This study provides estimations of wave conditions and the atmosphere stability that has an influence on wind wave. Results are also compared with data that have been recorded by a buoy in Caspian Sea (Neka region) during 1989. In the second part of this work we estimate non- dimensional energy and non-dimensional peak frequencies as a function of the non- dimensional fetch and Bulk Richardson numbers for the Persian Gulf (Busher region).This results also agree well with similar results for the Caspian Sea. The acquired relations can be used to compute the wind wave parameters. Also the results for the Persian Gulf show that the relationship of non-dimensional energy to as a function of wave age is independent of presence of swell. Finally the WAM model was run for the Persian Gulf during 3-8 September of 2002. The results show that swell on the Persian Gulf reduces the energy density of wind waves by up to 10%, but the growth rate at peak frequency is only reduced by up to 4%, and the spectral peak frequency is increased by only 1%.

The transport of nitric oxide in the upper atmosphere by planetary waves and the zonal mean circulation

U of I Only

In-situ based reanalysis of the global ocean temperature and salinity with ISAS: variability of the heat content and steric height

The In Situ Analysis System (ISAS) was developed to produce gridded fields of temperature and salinity that preserve as much as possible the time and space sampling capabilities of the Argo network of profiling floats. Since the first global re-analysis performed in 2009, the system has evolved and a careful delayed mode processing of the 2002-2012 dataset has been carried out using version 6 of ISAS and updating the statistics to produce the ISAS13 analysis. This last version is now implemented as the operational analysis tool at the Coriolis data centre. The robustness of the results with respect to the system evolution is explored through global quantities of climatological interest: the Ocean Heat Content and the Steric Height. Estimates of errors consistent with the methodology are computed. This study shows that building reliable statistics on the fields is fundamental to improve the monthly estimates and to determine the absolute error bars. The new mean fields and variances deduced from the ISAS13 re-analysis and dataset show significant changes relative to the previous ISAS estimates, in particular in the southern ocean, justifying the iterative procedure. During the decade covered by Argo, the intermediate waters appear warmer and saltier in the North Atlantic and fresher in the Southern Ocean than in WOA05 long term mean. At inter-annual scale, the impact of ENSO on the Ocean Heat Content and Steric Height is observed during the 2006-2007 and 2009-2010 events captured by the network.

Fukushima-derived radionuclides in sediments of the Japanese Pacific Ocean coast and various Japanese water samples (seawater, tap water, and coolant water of Fukushima Daiichi reactor unit 5)

We investigated Ocean sediments and seawater from inside the Fukushima exclusion zone and found radiocesium (134Cs and 137Cs) up to 800 Bq kg-1 as well as 90Sr up to 5.6 Bq kg-1. This is one of the first reports on radiostrontium in sea sediments from the Fukushima exclusion zone. Seawater exhibited contamination levels up to 5.3 Bq kg-1 radiocesium. Tap water from Tokyo from weeks after the accident exhibited detectable but harmless activities of radiocesium (well below the regulatory limit). Analysis of the Unit 5 reactor coolant (finding only 3H and even low 129I) leads to the conclusion that the purification techniques for reactor coolant employed at Fukushima Daiichi are very effective.

Inertial dissipation method applied to derive turbulent fluxed over the ocean during the Surface of the Ocean, Fluxes and Interactions with the Atmosphere Atlantic Stratocumulus Transition Experiment (SOFIA/ASTEX) and Structure des Echanges Mer-Atmosphere, Proprietes des Heterogeneites Oceaniques: Recherche Experimentale (SEMAPHORE) experiments with low to moderate wind speed

The transfer coefficients for momentum and heat have been determined for 10 m neutral wind speeds (U-10n) between 0 and 12 m/s using data from the Surface of the Ocean, Fluxes and Interactions with the Atmosphere (SOFIA) and Structure des Echanges Mer-Atmosphere, Proprietes des Heterogeneites Oceaniques: Recherche Experimentale (SEMAPHORE) experiments. The inertial dissipation method was applied to wind and pseudo virtual temperature spectra from a sonic anemometer, mounted on a platform (ship) which was moving through the turbulence held. Under unstable conditions the assumptions concerning the turbulent kinetic energy (TKE) budget appeared incorrect. Using a bulk estimate for the stability parameter, Z/L (where Z is the height and L is the Obukhov length), this resulted in anomalously low drag coefficients compared to neutral conditions. Determining Z/L iteratively, a low rate of convergence was achieved. It was concluded that the divergence of the turbulent transport of TKE was not negligible under unstable conditions. By minimizing the dependence of the calculated neutral drag coefficient on stability, this term was estimated at about -0.65Z/L. The resulting turbulent fluxes were then in close agreement with other studies at moderate wind speed. The drag and exchange coefficients for low wind speeds were found to be C-en x 10(3) = 2.79U(10n)(-1) + 0.66 (U-10n < 5.2 m/s), C-en x 10(3) = C-hn x 10(3) = 1.2 (U-10n greater than or equal to 5.2 m/s), and C-dn x 10(3) = 11.7U(10n)(-2) + 0.668 (U-10n < 5.5 m/s), which imply a rapid increase of the coefficient values as the wind decreased within the smooth flow regime. The frozen turbulence hypothesis and the assumptions of isotropy and an inertial subrange were found to remain valid at these low wind speeds for these shipboard measurements. Incorporation of a free convection parameterization had little effect.

Foraging behaviour, swimming performance and malformations of early stages of commercially important fishes under ocean acidification and warming

Early life stages of many marine organisms are being challenged by climate change, but little is known about their capacity to tolerate future ocean conditions. Here we investigated a comprehensive set of biological responses of larvae of two commercially important teleost fishes, Sparus aurata (gilthead seabream) and Argyrosomus regius (meagre), after exposure to future predictions of ocean warming (+4 °C) and acidification (ΔpH= 0.5). The combined effect of warming and hypercapnia elicited a decrease in the hatching success (by 26.4 and 14.3 % for S. aurata and A. regius, respectively) and larval survival (by half) in both species. The length for newly-hatched larvae was not significantly affected, but a significant effect of hypercapnia was found on larval growth. However, while S. aurata growth was reduced (24.8–36.4 % lower), A. regius growth slightly increased (3.2–12.9 % higher) under such condition. Under acidification, larvae of both species spent less time swimming, and displayed reduced attack and capture rates of prey. The impact of warming on these behavioural traits was opposite but less evident. While not studied in A. regius, the incidence of body malformations in S. aurata larvae increased significantly (more than tripled) under warmer and hypercapnic conditions. These morphological impairments and behavioural changes are expected to affect larval performance and recruitment success, and further influence the abundance of fish stocks and the population structure of these commercially important fish species. However, given the pace of ocean climate change, it is important not to forget that species may have the opportunity to acclimate and adapt.