Archive for the 'Science' Category

Assessing the impact of different carbonate system parameters on benthic foraminifera from controlled growth experiments

Insights into past marine carbon cycling and water mass properties can be obtained by means of geochemical proxies calibrated through controlled laboratory experiments with accurate seawater carbonate system (C-system) manipulations. Here, we explored the use of strontium/calcium ratio (Sr/Ca) of the calcite shells of benthic foraminifera as a potential seawater C-system proxy through a controlled growth experiment with two deep-sea species (Bulimina marginata and Cassidulina laevigata) and one intertidal species (Ammonia T6). To this aim, we used two experimental set-ups to decouple as much as possible the individual components of the carbonate system, i.e., changing pH at constant dissolved inorganic carbon (DIC) and changing DIC at constant pH. Four climatic chambers were used with different controlled concentrations of atmospheric pCO2 (180 ppm, 410 ppm, 1000 ppm, 1500 ppm). Our results demonstrated that pH did not influence the survival and growth of the three species. However, low DIC conditions (879 μmol kg−1) negatively affected B. marginata and C. laevigata through reduced growth, whereas no effect was observed for Ammonia T6. Our results also showed that Sr/Ca was positively correlated with total Alkalinity (TA), DIC and bicarbonate ion concentration ([HCO3]) for Ammonia T6 and B. marginata; i.e., DIC and/or [HCO3] were the main controlling factors. For these two species, the regression models were coherent with published data (existing so far only for Ammonia T6) and showed overall similar slopes but different intercepts, implying species-specific effects. Furthermore, the Sr/Ca – C-system relationship was not impacted by ontogenetic trends between chamber stages, which is a considerable advantage for paleo-applications. This applied particularly to Ammonia T6 that calcified many chambers compared to the two other species. However, no correlation with any of the C-system parameters was observed for Sr/Ca in C. laevigata. This might imply either a strong species-specific effect and/or a low tolerance to laboratory conditions leading to a physiological stress, thereby impacting the Sr incorporation into the calcite lattice of C. laevigata.

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Ocean acidification, warming and feeding impacts on biomineralization pathways and shell material properties of Magallana gigas and Mytilus spp.


  • Mytilus spp. source environmental carbon into the shell aragonite under low pH.
  • In Mytilus spp. biomineralization pathways differ between calcite and aragonite.
  • M. gigas carbon sourcing remains similar maintaining calcite growth.
  • M. gigas mantle δ15N is lower in low pH reflecting algae nitrogen uptake.
  • Calcite biomineralization pathway differs between the two species under low pH.


Molluscs are among the organisms affected by ocean acidification (OA), relying on carbon for shell biomineralization. Metabolic and environmental sourcing are two pathways potentially affected by OA, but the circumstances and patterns by which they are altered are poorly understood. From previous studies, mollusc shells grown under OA appear smaller in size, brittle and thinner, suggesting an important alteration in carbon sequestration. However, supplementary feeding experiments have shown promising results in offsetting the negative consequences of OA on shell growth. Our study compared carbon uptake by δ13C tracing and deposition into mantle tissue and shell layers in Magallana gigas and Mytilus species, two economically valuable and common species. After subjecting the species to 7.7 pH, +2 °C seawater, and enhanced feeding, both species maintain shell growth and metabolic pathways under OA without benefitting from extra feeding, thus, showing effective acclimation to rapid and short-term environmental change. Mytilus spp. increases metabolic carbon into the calcite and environmental sourcing of carbon into the shell aragonite in low pH and high temperature conditions. Low pH affects M. gigas mantle nitrogen isotopes maintaining growth. Calcite biomineralization pathway differs between the two species and suggests species-specific response to OA.

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The multi-generational effect of seawater acidification on larval development, reproduction, ingestion rate, and ATPase activity of Tigriopus japonicus Mori, 1938

Ocean acidification threatens marine organisms continuously. To ascertain if adaptation of marine species to ocean acidification enhanced over multiple generations, we studied the transgenerational effects of ocean acidification on the development, reproduction, ingestion rate, and ATPase activity of a copepod Tigriopus japonicus Mori, 1938. In the first mode, individuals were exposed to either one of the pH levels (8.1 (control), 7.7, 7.3) for five successive generations. In the second mode, each successive generation was exposed to a lower pH level (pH levels: 8.1, 7.9, 7.7, 7.5, 7.3). After prolonged exposure to a constant seawater acidification level, the capacity to adapt to the stress increased. However, when exposed to seawater of descending pH, the detrimental effects gradually increased. Energy allocated to development and reproduction was reduced although the ingestion rate continued to improve in successive generations. Therefore, ongoing ocean acidification might lower the energy transfer of copepods to higher trophic levels.

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Aquatic productivity under multiple stressors

Aquatic ecosystems are responsible for about 50% of global productivity. They mitigate climate change by taking up a substantial fraction of anthropogenically emitted CO2 and sink part of it into the deep ocean. Productivity is controlled by a number of environmental factors, such as water temperature, ocean acidification, nutrient availability, deoxygenation and exposure to solar UV radiation. Recent studies have revealed that these factors may interact to yield additive, synergistic or antagonistic effects. While ocean warming and deoxygenation are supposed to affect mitochondrial respiration oppositely, they can act synergistically to influence the migration of plankton and N2-fixation of diazotrophs. Ocean acidification, along with elevated pCO2, exhibits controversial effects on marine primary producers, resulting in negative impacts under high light and limited availability of nutrients. However, the acidic stress has been shown to exacerbate viral attacks on microalgae and to act synergistically with UV radiation to reduce the calcification of algal calcifiers. Elevated pCO2 in surface oceans is known to downregulate the CCMs (CO2 concentrating mechanisms) of phytoplankton, but deoxygenation is proposed to enhance CCMs by suppressing photorespiration. While most of the studies on climate-change drivers have been carried out under controlled conditions, field observations over long periods of time have been scarce. Mechanistic responses of phytoplankton to multiple drivers have been little documented due to the logistic difficulties to manipulate numerous replications for different treatments representative of the drivers. Nevertheless, future studies are expected to explore responses and involved mechanisms to multiple drivers in different regions, considering that regional chemical and physical environmental forcings modulate the effects of ocean global climate changes.

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Climate change effects on marine species across trophic levels

Climate change and anthropogenic activities are producing a range of new selection pressures, both abiotic and biotic, on marine organisms. While there are numerous studies that have investigated the response of individual marine organisms to climate change, few studies have focused on differences in organismal responses across trophic levels. Such trophic differences in response to climate change may disrupt ecological interactions and thereby threaten marine ecosystem function. In addition, predation is known as a strong driver that impacts individuals and populations. Despite this, we still do not have a comprehensive understanding of how different trophic levels respond to climate change stressors, predation and their combined effects in marine ecosystems.

The main focus of this thesis is to identify whether marine trophic levels respond differently to climatic stressors and predation. To explore these questions, I have used a combination of traditional mesocosm experiments, together with a statistical method called meta-analysis. I initiated the research by study the responses of marine gastropods at two trophic levels to ocean acidification and predation using long-term mesocosm experiments together with a gastropod-specific meta-analyses. I focused on the amount of phenotypic plasticity in morphological traits of snails when exposed to the two stressors. In order to generalise and test these assumptions among a greater number of marine taxa, I used the meta-analysis approach to investigate the effects of ocean acidification and warming, as well as their combined effects on four marine trophic levels. Finally, to study the individual and combined effects of ocean acidification and predation with respect to inducible defences, I again applied a mesocosm experiment and used blue mussels as a model species.

By using long-term mesocosm experiments and the gastropod-specific meta-analysis on marine gastropods from two trophic levels, I showed that these trophic levels varied in their responses to both ocean acidification and predation. Gastropods at lower trophic levels exhibited greater phenotypic plasticity against predation, while those from higher trophic levels showed stronger tolerance to ocean acidification. Next, by using a meta-analysis, including a large number of species and taxa, examining the effects of ocean acidification and warming, I revealed that top-predators and primary producers were most tolerant to ocean acidification compared to other trophic levels. Herbivores on the other hand, were the most vulnerable trophic level against abiotic stress. Again, using the meta-analysis approach, but this time incorporating only factorial experimental data that included the interactive effects of ocean acidification and ocean warming, I showed that higher trophic levels again were the most tolerant trophic level, and herbivores being most sensitive, with respect to the combined effect of the two stressors. Contrary to previous discussions in the literature concerning multiple climate-related stressors, antagonistic and additive effects occurred most frequently, while synergistic effects were less common and which decreased with increasing trophic rank. Finally, by conducting a fully-factorial experiment using blue mussels, I found that mussels with previous experience contact with predator has developed greater inducible defences than ones without previous experience. However, levels of ocean acidification may mask predator cues, or obstruct shell material, and consequently disrupt blue mussels inducible defence from crab predation.

In summary, marine trophic levels respond differently to both biotic and climatic stressors. Higher trophic levels, together with primary producers, were often more robust against abiotic stress and may therefore be better prepared for future oceans compare species from lower trophic levels. These results may provide vital information for: implementing effective climate change mitigation, to understand which stressors to act on, and when and where to intervene for prioritizing conservation actions.

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Early detection of anthropogenic climate change signals in the ocean interior

Robust detection of anthropogenic climate change is crucial to: (i) improve our understanding of Earth system responses to external forcing, (ii) reduce uncertainty in future climate projections, and (iii) develop efficient mitigation and adaptation plans. Here, we use Earth system model projections to establish the detection timescales of anthropogenic signals in the global ocean through analyzing temperature, salinity, oxygen, and pH evolution from surface to 2000 m depths. For most variables, anthropogenic changes emerge earlier in the interior ocean than at the surface, due to the lower background variability at depth. Acidification is detectable earliest, followed by warming and oxygen changes in the subsurface tropical Atlantic. Temperature and salinity changes in the subsurface tropical and subtropical North Atlantic are shown to be early indicators for a slowdown of the Atlantic Meridional Overturning Circulation. Even under mitigated scenarios, inner ocean anthropogenic signals are projected to emerge within the next few decades. This is because they originate from existing surface changes that are now propagating into the interior. In addition to the tropical Atlantic, our study calls for establishment of long-term interior monitoring systems in the Southern Ocean and North Atlantic in order to elucidate how spatially heterogeneous anthropogenic signals propagate into the interior and impact marine ecosystems and biogeochemistry.

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The marine carbonate system variability in high meltwater season (Spitsbergen Fjords, Svalbard)


  • Spatial variability in hydrography and carbonate chemistry were investigated.
  • Lack of clear relationship between alkalinity and salinity in the surface water.
  • Effect of alkalinity fluxes from sediments on the bottom water was insignificant.
  • Freshening of the surface water reduces significantly saturation state of aragonite.


The spatial variability in hydrography (salinity and temperature) and carbonate chemistry (alkalinity – AT, total inorganic carbon concentration – CT, pH, CO2 partial pressure – pCO2, and the saturation state of aragonite – ΩAr) in high meltwater season (summer) was investigated in four Spitsbergen fjords – Krossfjorden, Kongsfjorden, Isfjorden, and Hornsund. It was found that the differences in hydrology entail spatial changes in the CO2 system structure. AT decline with decreasing salinity was evident, hence it is clear that freshwater input generally has a diluting effect and lowers AT in the surface waters of the Spitsbergen fjords. Significant surface water AT variability (1889–2261 µmol kg−1) reveals the complexity of the fjords’ systems with multiple freshwater sources having different alkalinity end-member characteristics and identifies the mean AT freshwater end-member of 595 ± 84 µmol kg−1 for the entire region. The effect of AT fluxes from sediments on the bottom water was rather insignificant, despite high AT values (2288–2666 μmol kg−1) observed in the pore waters. Low pCO2 results in surface water (200–295 μatm) points to intensive biological production, which can strongly affect the CT values, however, is less important for shaping alkalinity. It has also been shown that the freshening of the surface water in the fjords reduces significantly ΩAr (an increase in freshwater fraction contribution by 1% causes a decrease in ΩAr by 0.022). Although during the polar day, due to low pCO2, ΩAr values are still rather far from 1 (they ranged from 1.4 to 2.5), during polar night, when pCO2 values are much higher, ΩAr may drop markedly. This study highlights that the use of salinity to estimate the potential alkalinity can carry a high uncertainty, while good recognition of the surface water AT variability and its freshwater end-members is key to predict marine CO2 system changes along with the ongoing freshening of fjords waters due to climate warming.

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Meridional variability in multi-decadal trends of dissolved inorganic carbon in surface seawater of the western North Pacific along the 165°E line


Multi-decadal trends of the total dissolved inorganic carbon (DIC) concentrations and consequent ocean acidification in surface seawater were investigated on the basis of data from shipboard measurements conducted since 1996 along the 165°E repeat line in the western North Pacific. The observed trends exhibited clear meridional variabilities, with higher rates in the subtropical and tropical zones and lower rates in the subarctic zone, with a DIC range from +0.09 ± 0.14 to +1.64 ± 0.16 μmol kg−1 yr−1 and pH range from −0.0023 ± 0.0034 to −0.0281 ± 0.0059 decade−1. DIC and acidification trends were consistent with those expected from the atmospheric CO2 concentrations at nearly all latitudinal zones, but were significantly different at some latitudes. We attribute the significantly lower rates observed in the central western Subarctic Gyre and southern Subtropical Gyres primarily to the variabilities in upward DIC supply from the subsurface associated with the variability in ocean circulation. However, the higher rate observed to the south of the Kuroshio Extension appears to have been caused by the change in winter vertical mixing related to the change in its stable/unstable paths.

Key Points

  • Meridional variability was found in the trends of dissolved inorganic carbon in the surface layer along the 165°E repeat line
  • Extremely slow rates of increase observed in the subarctic and tropics are attributed to ocean circulation variability
  • A fast increase in the south of the Kuroshio Extension is likely associated with the variability in its path
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Ocean acidification drives gut microbiome changes linked to species-specific immune defence

Ocean acidification (OA) has important effects on the intrinsic phenotypic characteristics of many marine organisms. Concomitantly, OA can alter the extended phenotypes of these organisms by perturbing the structure and function of their associated microbiomes. It is unclear, however, the extent to which interactions between these levels of phenotypic change can modulate the capacity for resilience to OA. Here, we explored this theoretical framework assessing the influence of OA on intrinsic (immunological responses and energy reserve) and extrinsic (gut microbiome) phenotypic characteristics and the survival of important calcifiers, the edible oysters Crassostrea angulata and C. hongkongensis. After one-month exposure to experimental OA (pH 7.4) and control (pH 8.0) conditions, we found species-specific responses characterised by elevated stress (hemocyte apoptosis) and decreased survival in the coastal species (C. angulata) compared with the estuarine species (C. hongkongensis). Phagocytosis of hemocytes was not affected by OA but in vitro bacterial clearance capability decreased in both species. Gut microbial diversity decreased in C. angulata but not in C. hongkongensis. Overall, C. hongkongensis was capable of maintaining the homeostasis of the immune system and energy supply under OA. In contrast, C. angulata’s immune function was suppressed, and the energy reserve was imbalanced, which might be attributed to the declined microbial diversity and the functional loss of essential bacteria in the guts. This study highlights a species-specific response to OA determined by genetic background and local adaptation, shedding light on the understanding of host-microbiota-environment interactions in future coastal acidification.

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Deoxygenation, acidification and warming in Waquoit Bay, USA, and a shift to pelagic dominance

Coastal nutrient pollution, or eutrophication, is commonly linked to anthropogenic influences in terrestrial watersheds, where land-use changes often degrade water quality over time. Due to gradual changes, the management and monitoring of estuarine systems often lag environmental degradation. One example can be found at the Waquoit Bay National Estuarine Research Reserve, where we developed an analysis framework to standardize and analyze long-term trends in water quality and submerged vegetation data from monitoring programs that began in the 1990s. These programs started after the nearly complete loss of historically extensive Zostera marina (eelgrass) meadows throughout the estuary. Recently, eelgrass only persisted in small, undeveloped sub-embayments of the estuary, with conservative declines of over 97% in areal coverage. Over the past 2 decades, the average deoxygenation, acidification, and warming were −24.7 µmol O2 kg−1 (−11%), 0.006 µmol H+ kg−1 (+ 34%), and 1.0 °C (+ 4%), respectively. Along with the loss of eelgrass, there was also a decline in macroalgal biomass over 3 decades, resulting in a system dominated by pelagic metabolism, indicated by a 71% increase in water column chlorophyll a concentrations since 2009. This recent increase in phytoplankton biomass, which is highly mobile and transported throughout the estuary by tides, has resulted in recent degradation of isolated embayments despite their lower nutrient loads. This shift toward pelagic dominance in Waquoit Bay may indicate that other eutrophic and warming estuaries may also shift toward pelagic dominance in the future, as the Northeastern US is one of the fastest warming regions across the world.

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Predicting effects of multiple interacting global change drivers across trophic levels

Global change encompasses many co-occurring anthropogenic drivers, which can act synergistically or antagonistically on ecological systems. Predicting how different global change drivers simultaneously contribute to observed biodiversity change is a key challenge for ecology and conservation. However, we lack the mechanistic understanding of how multiple global change drivers influence the vital rates of multiple interacting species. We propose that reaction norms, the relationships between a driver and vital rates like growth, mortality, and consumption, provide insights to the underlying mechanisms of community responses to multiple drivers. Understanding how multiple drivers interact to affect demographic rates using a reaction-norm perspective can improve our ability to make predictions of interactions at higher levels of organization—that is, community and food web. Building on the framework of consumer–resource interactions and widely studied thermal performance curves, we illustrate how joint driver impacts can be scaled up from the population to the community level. A simple proof-of-concept model demonstrates how reaction norms of vital rates predict the prevalence of driver interactions at the community level. A literature search suggests that our proposed approach is not yet used in multiple driver research. We outline how realistic response surfaces (i.e., multidimensional reaction norms) can be inferred by parametric and nonparametric approaches. Response surfaces have the potential to strengthen our understanding of how multiple drivers affect communities as well as improve our ability to predict when interactive effects emerge, two of the major challenges of ecology today.

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Understanding the dynamic response of Durafet-based sensors: a case study from the Murderkill Estuary-Delaware Bay system (Delaware, USA)


  • A SeapHOx sensor package was deployed in a dynamic estuarine environment.
  • The responses of the Durafet’s internal and external reference electrodes were assessed.
  • Previously unreported dynamic errors in their temperature and salinity responses were characterized.
  • A dynamic sensor response correction for the external reference electrode was developed.


The use of Durafet-based sensors has proliferated in recent years, but their performance in estuarine waters (salinity < 20) where rapid changes in temperature and salinity are frequently observed requires further scrutiny. Here, the responses of the Honeywell Durafet and its internal (pHINT) and external (pHEXT) reference electrodes integrated into a SeapHOx sensor at the confluence of the Murderkill Estuary and Delaware Bay (Delaware, USA) were assessed over extensive ranges of temperature (1.34–32.27°C), salinity (1.17–29.82), and rates of temperature (dT/dt; −1.46 to +1.53°C (0.5 h)−1) and salinity (dSalt/dt; −3.55 to +11.09 (0.5 h)−1) change. Empirical analyses indicated dynamic errors in the temperature and salinity responses of the internal and external reference electrodes, respectively, driven by tidal mixing were introduced into our pH time-series. These dynamic errors drove large anomalies between pHINT and pHEXT (denoted ΔpHINT−EXT) that reached >±0.8 pH in winter when the lowest temperatures and maximum tidal salinity variability occurred and >±0.15 pH in summer when the highest temperatures and minimum tidal salinity variability occurred. The ΔpHINT−EXT anomalies demonstrated a clear linear relationship with dSalt/dt thereby making dSalt/dt the strongest limiting factor of reference electrode response in our application. A dynamic sensor response correction for the external reference electrode (solid-state chloiride ion-selective electrode, Cl-ISE) was also developed and applied in the voltage domain. This correction reduced winter and summer ΔpHINT−EXT anomaly ranges by >40% and 68.7%, respectively. Summer anomalies were notably reduced to <±0.04 pH across all measurements. Further, this correction also removed the first-order salinity dependence of these anomalies. Consequently, dynamic errors in reference electrode response cannot be ignored and must be considered in future experimental designs. Further work to better understand the dynamic temperature and salinity responses of both reference electrodes is underway. Ultimately, we hope this work will stimulate further discussion around the role and treatment of large ΔpHINT−EXT anomalies as a part of future data quality control and data reporting as well as the dynamic errors in reference electrode response that drive them in the context of Sensor Best Practices.

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Oceans and the changing climate

Increasing levels of atmospheric greenhouse gases are producing changes in the world’s oceans and coastal environments, such as increasing sea surface temperatures, ocean acidification, and rising sea levels. This chapter explores the human dimensions of three climate change impacts: (1) rising sea levels, measures to adapt, and the potential displacement of persons from eroding, low-lying coastal areas; (2) the migration of fish stocks to new habitats resulting from increasing seawater temperatures; and (3) the degradation of coral reefs and impacts to shellfish resulting from ocean acidification. With sea level rise, millions of people in low-lying coastal cities and small island developing states must adapt or be displaced, and some will become climate refugees. In the case of fisheries, distributions of some fish stocks are already changing because of increasing ocean temperatures. These shifts have great implications for both fishers and managers of marine resources. Finally, rising atmospheric concentrations of CO2 that lower the ocean’s pH make it more difficult for corals and shellfish to precipitate the calcium carbonate that forms their exoskeletons and shells, affecting users of tropical coral reef ecosystems as well as the shellfish aquaculture industry.

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Wanted dead or alive: skeletal structure alteration of cold-water coral Desmophyllum pertusum (Lophelia pertusa) from anthropogenic stressors

Ocean acidification (OA) has provoked changes in the carbonate saturation state that may alter the formation and structural biomineralisation of calcium carbonate exoskeletons for marine organisms. Biomineral production in organisms such as cold-water corals (CWC) rely on available carbonate in the water column and the ability of the organism to sequester ions from seawater or nutrients for the formation and growth of a skeletal structure. As an important habitat structuring species, it is essential to examine the impact that anthropogenic stressors (i.e., OA and rising seawater temperatures) have on living corals and the structural properties of dead coral skeletons; these are important contributors to the entire reef structure and the stability of CWC mounds. In this study, dead coral skeletons in seawater were exposed to various levels of pCO2 and different temperatures over a 12-month period. Nanoindentation was subsequently conducted to assess the structural properties of coral samples’ elasticity (E) and hardness (H), whereas the amount of dissolution was assessed through scanning electron microscopy. Overall, CWC samples exposed to elevated pCO2 and temperature show changes in properties which leave them more susceptible to breakage and may in turn negatively impact the formation and stability of CWC mound development.

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Ocean acidification-mediated food chain transfer of polonium between primary producers and consumers

Phytoplankton and zooplankton are key marine components that play an important role in metal distribution through a food web transfer. An increased phytoplankton concentration as a result of ocean acidification and warming are well-established, along with the fact that phytoplankton biomagnify 210Po by 3–4 orders of magnitude compared to the seawater concentration. This experimental study is carried out to better understand the transfer of polonium between primary producers and consumers. The experimental produced data highlight the complex interaction between the polonium concentration in zooplankton food, i.e. phytoplankton, its excretion via defecated fecal pellets, and its bioaccumulation at ambient seawater pH and a lower pH of 7.7, typical of ocean acidification scenarios in the open ocean. The mass of copepods recovered was 11% less: 7.7 pH compared to 8.2. The effects of copepod species (n = 3), microalgae species (n = 3), pH (n = 2), and time (n = 4) on the polonium activity in the fecal pellets (expressed as % of the total activity introduced through feeding) was tested using an ANOVA 4. With the exception of time (model: F20, 215 = 176.84, p < 0.001; time: F3 = 1.76, p = 0.16), all tested parameters had an impact on the polonium activity (copepod species: F2 = 169.15, p < 0.0001; algae species: F2 = 10.21, p < 0.0001; pH: F1 = 9.85, p = 0.002) with complex interactions (copepod x algae: F2 = 19.48, p < 0.0001; copepod x pH: F2 = 10.54, p < 0.0001; algae x pH: F2 = 4.87, p = 0.009). The experimental data underpin the hypothesis that metal bioavailability and bioaccumulation will be enhanced in secondary consumers such as crustacean zooplankton due to ocean acidification.

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Climate change impacts on eastern boundary upwelling systems

The world’s eastern boundary upwelling systems (EBUSs) contribute disproportionately to global ocean productivity and provide critical ecosystem services to human society. The impact of climate change on EBUSs and the ecosystems they support is thus a subject of considerable interest. Here, we review hypotheses of climate-driven change in the physics, biogeochemistry, and ecology of EBUSs; describe observed changes over recent decades; and present projected changes over the twenty-first century. Similarities in historical and projected change among EBUSs include a trend toward upwelling intensification in poleward regions, mitigatedwarming in near-coastal regions where upwelling intensifies, and enhanced water-column stratification and a shoaling mixed layer. However, there remains significant uncertainty in how EBUSs will evolve with climate change, particularly in how the sometimes competing changes in upwelling intensity, source-water chemistry, and stratification will affect productivity and ecosystem structure. We summarize the commonalities and differences in historical and projected change in EBUSs and conclude with an assessment of key remaining uncertainties and questions. Future studies will need to address these questions to better understand, project, and adapt to climate-driven changes in EBUSs.

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Enormously enhanced particulate organic carbon and nitrogen production by elevated CO2 and moderate aluminum enrichment in the coccolithophore Emiliania huxleyi

Aluminum (Al) is abundant and ubiquitous in the environment. However, little information is available on its effects on photosynthetic microbes in alkaline seawater. Thus, we investigated the physiological performance in the most cosmopolitan coccolithophorid, viz., Emiliania huxleyi, grown under low (410 µatm) and high (1000 µatm) CO2 levels in seawater having none (0 nM, NAl), low (0.2 µM, LAl) and high (2 µM, HAl) Al concentrations. Under low CO2 conditions, the specific growth rate showed no significant difference between the NAl and LAl treatments, which was higher than the HAL treatment. Elevated CO2 inhibited the growth rate in the NAl and LAl cultures but did not affect the HAl cultures. The addition of Al had no effects on (LAl) or slightly elevated (HAl) the particulate organic carbon (POC) production rate under low CO2 conditions. With increasing CO2 concentration, the production rate of POC was enhanced by 55.3 % during the NAl treatment and further increased by 22.3 % by adding 0.2 µM Al. The responses of particulate organic nitrogen (PON) production rate, cellular POC, and PON contents to the different treatments revealed the same pattern as those of the POC production rate. The particulate inorganic carbon (PIC) production rate and PIC/POC ratio were not affected by Al under low CO2 conditions. They were significantly decreased by elevated CO2 in the LAl and HAl cultures. Our results indicate that high CO2 could increase carbon export to ocean depths by elevating the efficiency of the biological pump at low Al levels occurring in natural seawater (0.2 μM), with potentially significant implications for the carbon cycle of the ocean under accelerating anthropogenic influences.

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Effects of elevated CO2 on metabolic rate and nitrogenous waste handling in the early life stages of yellowfin tuna (Thunnus albacares)

Graphical abstract


  • Little is known about how tuna species will respond to ocean acidification (OA).
  • CO2 altered nitrogenous waste excretion and metabolic rate in yolk sac larvae.
  • CO2 did not change yolk sac depletion in embryos.
  • CO2 did not alter nitrogen accumulation in yellowfin tuna.
  • Yellowfin tuna were more robust to CO2 than predicted.


Ocean acidification is predicted to have a wide range of impacts on fish, but there has been little focus on broad-ranging pelagic fish species. Early life stages of fish are thought to be particularly susceptible to CO2 exposure, since acid-base regulatory faculties may not be fully developed. We obtained yellowfin tuna (Thunnus albacares) from a captive spawning broodstock population and exposed them to control or 1900 μatm CO2 through the first three days of development as embryos transitioned into yolk sac larvae. Metabolic rate, yolk sac depletion, and oil globule depletion were measured to assess overall energy usage. To determine if CO2 altered protein catabolism, tissue nitrogen content and nitrogenous waste excretion were quantified. CO2 exposure did not significantly impact embryonic metabolic rate, yolk sac depletion, or oil globule depletion, however, there was a significant decrease in metabolic rate at the latest measured yolk sac larval stage (36 h post fertilization). CO2-exposure led to a significant increase in nitrogenous waste excretion in larvae, but there were no differences in nitrogen tissue accumulation. Nitrogenous waste accumulated in embryos as they developed but decreased after hatch, coinciding with a large increase in nitrogenous waste excretion and increased metabolic rate in newly hatched larvae. Our results provide insight into how yellowfin tuna are impacted by increases in CO2 in early development, but more research with higher levels of replication is needed to better understand long-term impacts and acid-base regulatory mechanisms in this important pelagic fish.

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Alkalinity biases in CMIP6 Earth System Models and implications for simulated CO2 drawdown via artificial alkalinity enhancement

The partitioning of CO2 between atmosphere and ocean depends to a large degree not only on the amount of dissolved inorganic carbon (DIC) but also of alkalinity in the surface ocean. That is also why, in the context of negative emission approaches ocean alkalinity enhancement is discussed as one potential approach. Although alkalinity is thus an important variable of the marine carbonate system little knowledge exists how its representation in models compares with measurements. We evaluated the large-scale alkalinity distribution in 14 CMIP6 models against the observational data set GLODAPv2 and showed that most models as well as the multi-model-mean underestimate alkalinity at the surface and in the upper ocean, while overestimating alkalinity in the deeper ocean. The decomposition of the global mean alkalinity biases into contributions from physical processes (preformed alkalinity), remineralization, and carbonate formation and dissolution showed that the bias stemming from the physical redistribution of alkalinity is dominant. However, below the upper few hundred meters the bias from carbonate dissolution can become similarly important as physical biases, while the contribution from remineralization processes is negligible. This highlights the critical need for better understanding and quantification of processes driving calcium carbonate dissolution in microenvironments above the saturation horizons, and implementation of these processes into biogeochemical models.

For the application of the models to assess the potential of ocean alkalinity enhancement to increase ocean carbon uptake and counteract ocean acidification, a back-of-the-envelope calculation was conducted with each model’s global mean surface alkalinity and DIC as input parameters. We find that the degree of compensation of DIC and alkalinity biases at the surface is more important for the marine CO2 uptake capacity than the alkalinity biases themselves. The global mean surface alkalinity bias relative to GLODAPv2 in the different models ranges from -85 mmol kg-1 (-3.6 %) to +50 mmol kg-1 (+2.1 %) (mean: -25 mmol kg-1 or -1.1 %), while for DIC the relative bias ranges from -55 mmol kg-1 (-2.6 %) to 53 mmol kg-1 (+2.5 %) (mean: -13 mmol kg-1 or -0.6 %). Because of this partial compensation, all but two of the CMIP6 models evaluated here overestimate the Revelle factor at the surface and thus overestimate the CO2-draw-down after alkalinity addition by up to 13 % and pH increase by up to 7.2 %. This overestimate has to be taken into account when reporting on efficiencies of ocean alkalinity enhancement experiments using CMIP6 models.

Continue reading ‘Alkalinity biases in CMIP6 Earth System Models and implications for simulated CO2 drawdown via artificial alkalinity enhancement’

Effects of climate change on the Kenyan coral reef eco-system

The coral reef ecosystem is a natural habitat for many marine organisms that has high economic and tourist significance. Nonetheless, this ecosystem has very low tolerance to the effects of changes brought about by increasing sea surface temperatures and ocean acidification. This study sought to investigate the combined effect of rising sea surface temperatures and ocean acidification on the Kenyan coral reef ecosystem. This was achieved by determining the spatial-temporal variability of ocean acidification over the Kenyan coastline; and simulating the combined effect of sea surface temperature increases and ocean acidification on the coral reef ecosystem.

Historical (2000-2021) data on sea surface temperature (SSTs) was obtained from the National Oceanic and Atmospheric Administration (NOAA) and data on dissolved total carbon dioxide (TCO2) and pH from Global Ocean Data Analysis Project (GLODAP). Future (2022-2081) sea surface temperature and dissolved carbon dioxide data was downloaded from Coupled Model Intercomparison Project (CMIP6) experiment for two Shared Socioeconomic Pathways (SSPs) namely SSP2-4.5 and SSP5-8.5. Statistical, graphical and model simulations analyses were applied in the study to investigate the combined effect of increasing SST and ocean acidification on coral reef ecosystem over the Kenyan coastline.

Results indicate that mean sea surface temperature and dissolved carbon dioxide along the Kenyan coastline varied with seasons and had increased between the years 2000-2021. Trend tests of SSTs and TCO2 revealed a significant upward trend at 5% level of significance. Rising SSTs led to bleaching in coral reefs along this coastline whereas TCO2 led to reduced amount of carbonate ion concentration and reduced pH in the sea surface waters which affected the rates of calcification and survival of the coral reefs. The results of the Combined Mortality and Bleaching Output model simulation revealed that bleaching and ocean acidification had negatively affected the coral reef cover resulting in a decline of more than 30% of cover between 2000 and 2021. The results of the simulation also projected that the coral reef cover will continue to decline in the long-term by 52% under SSP2-4.5 and 63% under SSP5-8.5 if the trends in SSTs and TCO2 are maintained.

This study recommends collaborative implementation of climate change policies and practices by national and regional governments, communities and policy makers; enhanced efforts by coastal county governments in Kenya and research organisations to expound on scientific knowledge base while simultaneously implementing sustainable targeted solutions to ensure that the socio-economic benefits of the coral reef ecosystem are sustained.

Continue reading ‘Effects of climate change on the Kenyan coral reef eco-system’

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