Perspectives on Biological Research for CO<Subscript>2</Subscript> Ocean Sequestration

Feasibility studies recently suggest that sequestration of anthropogenic CO₂ in the deep ocean could help reduce the atmospheric CO₂ concentration. However, implementation of this strategy could have a significant environmental impact on marine organisms. This has highlighted the urgent need of further studies concerning the biological impact of CO₂ ocean sequestration. In this paper we summarize the recent literature reporting on the biological impact of CO₂ and discuss the research work required for the future. Although fundamental research of the effect of CO₂ on marine organisms before the practical consideration of CO₂ ocean sequestration was limited, laboratory and field studies concerning biological impacts have been increasing after the first international workshop in 1991 discussing CO₂ ocean sequestration. Acute impacts of CO₂ ocean sequestration could be determined by laboratory and field experiments and assessed by simulation models as described by the following papers in this section. On the other hand, chronic effects of CO₂ ocean sequestration, those directly related to the marine ecosystem, would be difficult to verify by means of experiments and to assess using ecosystem models. One of the practical solutions for this issue implies field experiments starting with controlled small scale and eventually to a large scale of CO₂ injection intended to determine ecosystem alteration. This revised version was published online in July 2006 with corrections to the Cover Date.     

A Eulerian-Eulerian Physical-Biological Impact Model of Zooplankton Injury Due to CO<Subscript>2</Subscript> Ocean Sequestration

To study the biological impacts of CO₂ ocean sequestration on floating marine organisms, a full Eulerian-Eulerian scheme model has been developed in a large-eddy simulation (LES) version using one-way coupling of the equations of seawater flow to the transport equations of the bio-scalar variables. Special attention was paid to deriving the transport equation, involving non-conservative scalars to describe the degree of injury to floating organisms due to the change in the pH environment resulting from CO₂ dissolution. The source terms of the transport equations of bio-scalar variables are based on experimental data on zooplankton activities affected by lower pH seawater, allowing construction of empirical sub-models of three kinds of floating marine organisms: Gaidius variabilis, Paraeuchaeta Birostrata, and Multi-organisms. An example is given to show the applicability of the model to the assessment of the biological impact of CO₂ sequestration in the ocean. Given an initial CO₂ droplet diameter of 8.0 mm and an injection rate of 1.0 kg/sec, the model simulation predicts that the zooplanktons lose approximately 90% of their activity when the lowest pH inside the plume decreases from 7.57 to 5.61. These injured zooplanktons then recovered gradually to their normal state within two hours due to dilution of the plume. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of Direct Ocean CO<Subscript>2</Subscript> Injection on Deep-Sea Meiofauna

Purposeful deep-sea carbon dioxide sequestration by direct injection of liquid CO₂ into the deep waters of the ocean has the potential to mitigate the rapid rise in atmospheric levels of greenhouse gases. One issue of concern for this carbon sequestration option is the impact of changes in seawater chemistry caused by CO₂ injection on deep-sea ecosystems. The effects of deep-sea carbon dioxide injection on infaunal deep-sea organisms were evaluated during a field experiment in 3600 m depth off California, in which liquid CO₂ was released on the seafloor. Exposure to the dissolution plume emanating from the liquid CO₂ resulted in high rates of mortality for flagellates, amoebae, and nematodes inhabiting sediments in close proximity to sites of CO₂ release. Results from this study indicate that large changes in seawater chemistry (i.e. pH reductions of ∼0.5–1.0 pH units) near CO₂ release sites will cause high mortality rates for nearby infaunal deep-sea communities. This revised version was published online in July 2006 with corrections to the Cover Date.     

Influence of Introduced CO<Subscript>2</Subscript> on Deep-Sea Metazoan Meiofauna

An experiment was performed to determine the effect of injected CO₂ on the deep-sea (3200 m) meiofaunal community in the Monterey Canyon. Approximately 20 L of liquid CO₂ was added to each of three cylindrical corrals (PVC rings pushed into the seabed) that were arranged in a triangular array 10 m on a side. After a 30-day period, sediment cores were collected within an area exposed to the dissolution plume emanating from the CO₂ pools and from a reference site approximately 40 m away; cores were also collected from within two of the CO₂ corrals. Sediment cores were sectioned into 0–5, 5–10, and 10–20 mm layers. Abundances of major groups (harpacticoid copepods, nematodes, nauplii, kinorhynchs, polychaetes, and total meiofauna) were determined for each layer. CO₂ exposure did not significantly influence the abundances or vertical distributions of any of the major taxa. However, other evidence suggests that abundance alone did not accurately reflect the effect of CO₂ on meiofauna. We argue that slow decomposition rates of meiofaunal carcasses can mask adverse effects of CO₂ and that longer experiments and/or careful examination of meiofaunal condition are needed to accurately evaluate CO₂ effects on deep-sea meiofaunal communities. This revised version was published online in July 2006 with corrections to the Cover Date.     

Sub-Lethal Effects of Elevated Concentration of CO<Subscript>2</Subscript> on Planktonic Copepods and Sea Urchins

Data concerning the effects of high CO₂ concentrations on marine organisms are essential for both predicting future impacts of the increasing atmospheric CO₂ concentration and assessing the effects of deep-sea CO₂sequestration. Here we review our recent studies evaluating the effects of elevated CO₂ concentrations in seawater on the mortality and egg production of the marine planktonic copepod, Acartia steueri, and on the fertilization rate and larval morphology of sea urchin embryos, Hemicentrotus pulcherrimus and Echinometra mathaei. Under conditions of +10,000 ppm CO₂ in seawater (pH 6.8), the egg production rates of copepods decreased significantly. The survival rates of adult copepods were not affected when reared under increased CO₂ for 8 days, however longer exposure times could have revealed toxic effects of elevated CO₂ concentrations. The fertilization rate of sea urchin eggs of both species decreased with increasing CO₂ concentration. Furthermore, the size of pluteus larvae decreased with increasing CO₂ concentration and malformed skeletogenesis was observed in both larvae. This suggests that calcification is affected by elevated CO₂ in the seawater. From these results, we conclude that increased CO₂ concentration in seawater will chronically affect several marine organisms and we discuss the effects of increased CO₂ on the marine carbon cycle and marine ecosystem. This revised version was published online in July 2006 with corrections to the Cover Date.     

Biological Impact of Elevated Ocean CO<Subscript>2</Subscript> Concentrations: Lessons from Animal Physiology and Earth History

CO₂ currently accumulating in the atmosphere permeates into ocean surface layers, where it may impact on marine animals in addition to effects caused by global warming. At the same time, several countries are developing scenarios for the disposal of anthropogenic CO₂ in the worlds' oceans, especially the deep sea. Elevated CO₂ partial pressures (hypercapnia) will affect the physiology of water breathing animals, a phenomenon also considered in recent discussions of a role for CO₂ in mass extinction events in earth history. Our current knowledge of CO₂ effects ranges from effects of hypercapnia on acid-base regulation, calcification and growth to influences on respiration, energy turnover and mode of metabolism. The present paper attempts to evaluate critical processes and the thresholds beyond which these effects may become detrimental. CO₂ elicits acidosis not only in the water, but also in tissues and body fluids. Despite compensatory accumulation of bicarbonate, acid-base parameters (pH, bicarbonate and CO₂ levels) and ion levels reach new steady-state values, with specific, long-term effects on metabolic functions. Even though such processes may not be detrimental, they are expected to affect long-term growth and reproduction and may thus be harmful at population and species levels. Sensitivity is maximal in ommastrephid squid, which are characterized by a high metabolic rate and extremely pH-sensitive blood oxygen transport. Acute sensitivity is interpreted to be less in fish with intracellular blood pigments and higher capacities to compensate for CO₂ induced acid-base disturbances than invertebrates. Virtually nothing is known about the degree to which deep-sea fishes are affected by short or long term hypercapnia. Sensitivity to CO₂ is hypothesized to be related to the organizational level of an animal, its energy requirements and mode of life. Long-term effects expected at population and species levels are in line with recent considerations of a detrimental role of CO₂ during mass extinctions in the earth's history. Future research is needed in this area to evaluate critical effects of the various CO₂ disposal scenarios. This revised version was published online in July 2006 with corrections to the Cover Date.     

Air-sea CO<Subscript>2</Subscript> flux by eddy covariance technique in the equatorial Indian Ocean

Direct measurements of the air-sea CO₂ flux by the eddy covariance technique were carried out in the equatorial Indian Ocean. The turbulent flux observation system was installed at the top of the foremast of the R/V MIRAI, thus minimizing dynamical and thermal effects of the ship body. During the turbulent flux runs around the two stations, the vessel was steered into the wind at constant speed. The power spectra of the temperature or water vapor density fluctuations followed the Kolmogorov −5/3 power law, although that of the CO₂ density fluctuation showed white noise in the high frequency range. However, the cospectrum of the vertical wind velocity and CO₂ density was well matched with those of the vertical velocity and temperature or water vapor density in this frequency range, and the CO₂ white noise did not influence the CO₂ flux. The raw CO₂ fluxes due to the turbulent transport showed a sink from the air to the ocean, and had almost the same value as the source CO₂ fluxes due to the mean vertical flow, corrected by the sensible and latent heat fluxes (called the Webb correction). The total CO₂ fluxes including the Webb correction terms showed a source from the ocean to the air, and were larger than the bulk CO₂ fluxes estimated using the gas transfer velocity by mass balance techniques.     

Effects of CO<Subscript>2</Subscript> Enrichment on Marine Phytoplankton

Rising atmospheric CO₂ and deliberate CO₂ sequestration in the ocean change seawater carbonate chemistry in a similar way, lowering seawater pH, carbonate ion concentration and carbonate saturation state and increasing dissolved CO₂ concentration. These changes affect marine plankton in various ways. On the organismal level, a moderate increase in CO₂ facilitates photosynthetic carbon fixation of some phytoplankton groups. It also enhances the release of dissolved carbohydrates, most notably during the decline of nutrient-limited phytoplankton blooms. A decrease in the carbonate saturation state represses biogenic calcification of the predominant marine calcifying organisms, foraminifera and coccolithophorids. On the ecosystem level these responses influence phytoplankton species composition and succession, favouring algal species which predominantly rely on CO₂ utilization. Increased phytoplankton exudation promotes particle aggregation and marine snow formation, enhancing the vertical flux of biogenic material. A decrease in calcification may affect the competitive advantage of calcifying organisms, with possible impacts on their distribution and abundance. On the biogeochemical level, biological responses to CO₂ enrichment and the related changes in carbonate chemistry can strongly alter the cycling of carbon and other bio-active elements in the ocean. Both decreasing calcification and enhanced carbon overproduction due to release of extracellular carbohydrates have the potential to increase the CO₂ storage capacity of the ocean. Although the significance of such biological responses to CO₂ enrichment becomes increasingly evident, our ability to make reliable predictions of their future developments and to quantify their potential ecological and biogeochemical impacts is still in its infancy. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of CO<Subscript>2</Subscript> on Marine Fish: Larvae and Adults

CO₂-enriched seawater was far more toxic to eggs and larvae of a marine fish, silver seabream, Pagrus major, than HCl-acidified seawater when tested at the same seawater pH. Data on the effects of acidified seawater can therefore not be used to estimate the toxicity of CO₂, as has been done in earlier studies. Ontogenetic changes in CO₂ tolerance of two marine bony fishes (Pag. major and Japanese sillago, Sillago japonica) showed a similar, characteristic pattern: the cleavage and juvenile stages were most susceptible, whereas the preflexion and flexion stages were much more tolerant to CO₂. Adult Japanese amberjack, Seriola quinqueradiata, and bastard halibut, Paralichthys olivaceus, died within 8 and 48 h, respectively, during exposure to seawater equilibrated with 5% CO₂. Only 20% of a cartilaginous fish, starspotted smooth-hound, Mustelus manazo, died at 7% CO₂ within 72 h. Arterial pH initially decreased but completely recovered within 1-24 h for Ser. quinqueradiata and Par. olivaceus at 1 and 3% CO₂, but the recovery was slower and complete only at 1% for M. manazo. During exposure to 5% CO₂, Par. olivaceus died after arterial pH had been completely restored. Exposure to 5% CO₂ rapidly depressed the cardiac output of Ser. quinqueradiata, while 1% CO₂ had no effect. Both levels of ambient CO₂ had no effect on blood O₂ levels. We tentatively conclude that cardiac failure is important in the mechanisms by which CO₂ kills fish. High CO₂ levels near injection points during CO₂ ocean sequestration are likely to have acute deleterious effects on both larvae and adults of marine fishes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Ventilation time and anthropogenic CO<Subscript>2</Subscript> in the Bering Sea and the Arctic Ocean based on carbon tetrachloride measurements

The Arctic Ocean is connected to the Pacific by the Bering Sea and the Bering Strait. During the 4th Chinese National Arctic Research Expedition, measurements of carbon tetrachloride (CCl₄) were used to estimate ventilation time-scales and anthropogenic CO₂ (C_ant) concentrations in the Arctic Ocean and Bering Sea based on the transit time distribution method. The profile distribution showed that there was a high-CCl₄ tongue entering through the Canada Basin in the intermediate layer (27.6?<?σ_θ?<?28), at latitudes between 78 and 85°N, which may be related to the inflow of Atlantic water. Between stations B09 and B10, upwelling appeared to occur near the continental slope in the Bering Sea. The ventilation time scales (mean ages) for deep and bottom water in the Arctic Ocean (~?230–380 years) were shorter than in the Bering Sea (~?430–970 years). Higher mean ages show that ventilation processes are weaker in the intermediate water of the Bering Sea than in the Arctic Ocean. The mean C_ant column inventory in the upper 4000 m was higher (60–82 mol m^?2) in the Arctic Ocean compared to the Bering Sea (35–48 mol m^?2).     

Features of coastal upwelling regions that determine net air-sea CO<Subscript>2</Subscript> flux

The influence of the coastal ocean on global net annual air-sea CO₂ fluxes remains uncertain. However, it is well known that air-sea pCO₂ disequilibria can be large (ocean pCO₂ ranging from ∼400 μatm above atmospheric saturation to ∼250 μatm below) in eastern boundary currents, and it has been hypothesized that these regions may be an appreciable net carbon sink. In addition it has been shown that the high productivity in these regions (responsible for the exceptionally low surface pCO₂) can cause nutrients and inorganic carbon to become more concentrated in the lower layer of the water column over the shelf relative to adjacent open ocean waters of the same density. This paper explores the potential role of the winter season in determining the net annual CO₂ flux in temperate zone eastern boundary currents, using the results from a box model. The model is parameterized and forced to represent the northernmost part of the upwelling region on the North American Pacific coast. Model results are compared to the few summer data that exist in that region. The model is also used to determine the effect that upwelling and downwelling strength have on the net annual CO₂ flux. Results show that downwelling may play an important role in limiting the amount of CO₂ outgassing that occurs during winter. Finally data from three distinct regions on the Pacific coast are compared to highlight the importance of upwelling and downwelling strength in determining carbon fluxes in eastern boundary currents and to suggest that other features, such as shelf width, are likely to be important.     

Effects of seawater acidification by ocean CO<Subscript>2</Subscript> sequestration on bathypelagic prokaryote activities

We investigated the effects of seawater acidification induced by ocean CO₂ sequestration on bathypelagic prokaryotes. We simulated acidification conditions by bubbling high-CO₂ air or adding chemical buffer solutions to seawater samples in order to examine changes in total cell counts, heterotrophic production rate, direct viable cell count, and relative abundance of Bacteria and Archaea. Considerable suppression of prokaryotic activities was observed at pH 7.0 or lower, especially in samples enriched with organic matter. The relative abundance of Archaea increased with increasing CO₂ concentration. We found that seawater acidification can potentially alter heterotrophic activities and community structure of bathypelagic prokaryotes.     

Uptake mechanism of anthropogenic CO<Subscript>2</Subscript> in the Kuroshio Extension region in an ocean general circulation model

The uptake mechanism of anthropogenic CO₂ in the Kuroshio Extension is examined by a Lagrangian approach using a biogeochemical model embedded in an ocean general circulation model. It is found that the uptake of anthropogenic CO₂ is caused mainly by the increase of pCO₂ dependency of seawater on temperature, which is caused by greater dissolved inorganic carbon concentration in the modern state than in the pre-industrial state. In contrast with the view of previous studies, the effect of the vertical entrainment, which brings waters that last contacted the atmosphere with the past lower CO₂ concentration, is comparatively small. Winter uptake of anthropogenic CO₂ increases with the rise of the atmospheric CO₂ level, while summer uptake is relatively stable, resulting in a larger seasonal cycle of the uptake. This increase is significant, especially in the Kuroshio Extension region. It is newly suggested that this increase in the Kuroshio Extension region is largely caused by the combined effects of the increased pCO₂ dependency of the sea water on the temperature and the seasonal difference in cooling.     

Influence of export rain ratio changes on atmospheric CO<Subscript>2</Subscript> and sedimentary calcite preservation

The responses of atmospheric pCO₂ and sediment calcite content to changes in the export rain ratio of calcium carbonate to organic carbon are examined using a diffusion-advection ocean biogeochemical model coupled to a one-dimensional sediment geochemistry model. Our model shows that a 25% reduction in rain ratio decreases atmospheric pCO₂ by 59 ppm. This is caused by alkalinity redistribution by a weakened carbonate pump and an alkalinity increase in the whole ocean via carbonate compensation with decreasing calcite burial. The steady state responses of sedimentary calcite content and calcite preservation efficiency are rather insensitive to the deepening of the saturation horizon of 1.9 km. This insensitivity is a result of the reduced deposition flux that decreases calcite burial, counteracting the saturation horizon deepening that increases calcite burial. However, in the first 10,000 years the effect of reduced calcite deposition on the burial change is more prominent; while after 10,000 years, the effect of saturation horizon deepening is more dominant. The lowering of sediment calcite content for the first 10,000 years is effectively decoupled from the 1.9 km downward shift of the saturation horizon. Our results are in part a consequence of the more dominant role that respiration CO₂ plays in sediment calcite dissolution over bottom water chemistry in our control run and support the decoupling of calcite lysocline depth and saturation horizon shifts, as suggested originally by Archer and Maier-Reimer (1994) and Archer et al. (2000).     

CO<Subscript>2</Subscript> Flux Measurements over the Sea Surface by Eddy Correlation and Aerodynamic Techniques

The measurements of the vertical transport of CO₂ were carried out over the Sea of Japan using the specially designed pier of Kyoto University on September 20 to 22, 2000. CO₂ fluxes were measured by the eddy correlation and aerodynamic techniques. Both techniques showed comparable CO₂ fluxes during sea breeze conditions: −0.001 to −0.08 mg m⁻²s⁻¹ with the mean of −0.05 mg m⁻²s⁻¹. This means that the measuring site satisfies the fetch requirement for meteorological observations under sea breeze conditions. Moreover, the eddy diffusivity coefficient used in the aerodynamic technique is found to be consistent with the coefficient used in the eddy correlation technique. The present result leads us to conclude that the aerodynamic technique may be applicable to underway CO₂ flux measurements over the ocean and may be used in place of the bulk technique. The important point is the need to maintain a measuring accuracy of CO₂ concentration difference of the order of 0.1 ppmv on the research vessels or the buoys.     

Seasonal variability of surface and internal M<Subscript>2</Subscript> tides in the Arctic Ocean

The modeling results of surface and internal M₂ tides for summer and winter periods in the Arctic Ocean (AO) are presented. We employed a modified version of the three-dimensional finite-element hydrothermodynamic model QUODDY-4 differing from the original model by using a rotated (instead of spherical) coordinate system and by considering the equilibrium-tide effects. It has been shown that the modeling results for the surface tide differs little from the results obtained earlier by other authors. According to these results, the amplitudes of internal tidal waves (ITWs) in the AO are significantly lower than in other oceans and the ITWs proper have the character of trapped waves. Their source of generation is located at the continental slope northwest of the New Siberian Islands. Our results are consistent with the fields of average (over a tidal cycle) and integral (by depth) densities of baroclinic tidal energy, the maximum baroclinic tidal velocity, and the coefficient of diapycnic mixing. The local rate of baroclinic tidal energy dissipation at the AO ridges increases as it approaches the bottom, as was observed on Mid-Atlantic and Hawaii ridges (but merely within the bottom boundary layer) and is two to three orders of magnitude lower than in other oceans. The ITW degeneration scale in the AO is several hundreds of kilometers in summer and winter, remaining within the range of its values between 100 and 1000 km in mid- and low-latitude oceans. In both seasons, the integral (over the AO area) rate of baroclinic tidal energy dissipation is two orders of magnitude lower than the global estimate (2.5 × 10¹² W).     

Physiological Responses of the Mediterranean Subtidal Alga <Emphasis Type="Italic">Peyssonnelia squamaria</Emphasis> to Elevated CO<Subscript>2</Subscript>

The ecological consequences of ocean acidification are unclear due to varying physiological properties of macroalgae and species-specific responses. Therefore, in the present study, we used a laboratory culture experiment to analyse the eco-physiological responses of the Mediterranean subtidal red alga Peyssonnelia squamaria to CO₂-induced lower pH. Our results showed an increase in the photosynthetic performance and growth rate of P. squamaria, despite the reduction in CaCO₃ content in the low pH treatment. According to our results, we believe that samples exposed to elevated CO₂ could be regulated own nitrogen metabolism to support increased growth rate and it may be down-regulated nitrate uptake. As a result, we hypothesize that P. squamaria may benefit from ocean acidification.     

The spatial distribution of surface <Emphasis Type="Italic">f</Emphasis>CO<Subscript>2</Subscript> in the Southwestern East Sea/Japan sea during summer 2005

Fugacity of CO₂ (fCO₂), temperature, salinity, nutrients, and chlorophyll-a were measured in the surface waters of southwestern East Sea/Japan Sea in July 2005. Surface waters were divided into three waters based on hydrographic characteristics: the water with moderate sea surface temperature (SST) and high sea surface salinity (SSS) located east of the front (East water); the water with high SST and moderate SSS located west of the front (West water); and the water with low SST and SSS located in the middle part of the study area (Middle water). High fCO₂ larger than 420 μatm were found in the West water. In the Middle water, CO₂ was undersaturated with respect to the atmosphere, with values between 246 and 380 μatm. Moderate fCO₂ values ranging from 370 to 420 μatm were observed in the East water. For the East and West waters, estimates of temperature dependency of fCO₂ (12.6 and 15.1 μatm °C⁻¹, respectively) were rather similar to a theoretical value, indicating that SST is likely to be a major factor controlling the surface fCO₂ distribution in these two regions. In the Middle water, however, the estimated temperature dependence was somewhat lower than the theoretical value, and relatively high concentrations of surface chlorophyll-a coincided with the low surface fCO₂, implying that biological uptake may considerably affect the fCO₂ distribution. The net sea-to-air CO₂ flux of the study area was estimated to be 0.30±4.81 mmol m⁻² day⁻¹ in summer, 2005.     

Comparing data obtained from ground-based measurements of the total contents of O<Subscript>3</Subscript>, HNO<Subscript>3</Subscript>,HCl,and NO<Subscript>2</Subscript> and from their numerical simulation

Chemistry climate models of the gas composition of the atmosphere make it possible to simulate both space and time variations in atmospheric trace-gas components (TGCs) and predict their changes. Both verification and improvement of such models on the basis of a comparison with experimental data are of great importance. Data obtained from the 2009–2012 ground-based spectrometric measurements of the total contents (TCs) of a number of TGCs (ozone, HNO₃, HCl, and NO₂) in the atmosphere over the St. Petersburg region (Petergof station, St. Petersburg State University) have been compared to analogous EMAC model data. Both daily and monthly means of their TCs for this period have been analyzed in detail. The seasonal dependences of the TCs of the gases under study are shown to be adequately reproduced by the EMAC model. At the same time, a number of disagreements (including systematic ones) have been revealed between model and measurement data. Thus, for example, the EMAC model underestimates the TCs of NO₂, HCl, and HNO₃, when compared to measurement data, on average, by 14, 22, and 35%, respectively. However, the TC of ozone is overestimated by the EMAC model (on average, by 12%) when compared to measurement data. In order to reveal the reasons for such disagreements between simulated and measured data on the TCs of TGCs, it is necessary to continue studies on comparisons of the contents of TGCs in different atmospheric layers.     

The Interactive Effects of Elevated CO<Subscript>2</Subscript> and Ammonium Enrichment on the Physiological Performances of <Emphasis Type="Italic">Saccharina japonica</Emphasis> (Laminariales,Phaeophyta)

Environmental challenges such as ocean acidification and eutrophication influence the physiology of kelp species. We investigated their interactive effects on Saccharina japonica (Laminariales, Phaeophyta) under two pH conditions [Low, 7.50; High (control), 8.10] and three NH ₄ ⁺ concentrations (Low, 4; Medium, 60; High, 120 μM). The degree of variation of pH values in the culture medium and inhibition rate of photosynthetic oxygen evolution by acetazolamide were affected by pH treatments. Relative growth rates, carbon, nitrogen, and the C:N ratio in tissue samples were influenced by higher concentrations of NH ₄ ⁺ . Rates of photosynthetic oxygen evolution were enhanced under elevated CO₂ or NH ₄ ⁺ conditions, independently, but these two factors did not show an interactive effect. However, rates of NH ₄ ⁺ uptake were influenced by the interactive effect of increased CO₂ under elevated NH ₄ ⁺ treatment. Although ocean acidification and eutrophication states had an impact on physiological performance, chlorophyll fluorescence was not affected by those conditions. Our results indicated that the physiological reactions by this alga were influenced to some extent by a rise in the levels of CO₂ and NH ₄ ⁺ . Therefore, we expect that the biomass accumulation of S. japonica may well increase under future scenarios of ocean acidification and eutrophication.