The world's shellfish are under threat as our oceans become more acidic

First paragraph: For the past few million years the world’s oceans have existed in a slightly alkaline state, with an average pH of 8.2. Now, with carbon emissions escalating, there is more CO₂ in the world’s atmosphere. This dissolves in the oceans, altering the chemistry of the seawater by lowering the pH and making it more acidic – up to 30% more in the past 200 years. This growing acidification of the oceans is becoming a serious problem for the production of shellfish around the world.https://theconversation.com/the-worlds-shellfish-are-under-threat-as-our-oceans-become-more-acidic-10386

The impact of environmental acidification on the microstructure and mechanical integrity of marine invertebrate skeletons

Ocean acidification (OA), from seawater uptake of anthropogenic CO2, has a suite of negative effects on the ability of marine invertebrates to produce and maintain their skeletons. Increased organism pCO2 causes hypercapnia, an energetically costly physiological stress. OA alters seawater carbonate chemistry, limiting the carbonate available to form the calcium carbonate (CaCO3) minerals used to build skeletons. The reduced saturation state of CaCO3 also causes corrosion of CaCO3 structures. Global change is also accelerating coastal acidification driven by land-run off (e.g. acid soil leachates, tannic acid). Building and maintaining marine biomaterials in the face of changing climate will depend on the balance between calcification and dissolution. Overall, in response to environmental acidification, many calcifiers produce less biomineral and so have smaller body size. Studies of skeleton development in echinoderms and molluscs across life stages show the stunting effect of OA. For corals, linear extension may be maintained, but at the expense of less dense biomineral. Conventional metrics used to quantify growth and calcification need to be augmented by characterisation of the changes to biomineral structure and mechanical integrity caused by environmental acidification. Scanning electron microscopy and microcomputed tomography of corals, tube worms and sea urchins exposed to experimental (laboratory) and natural (vents, coastal run off) acidification show a less dense biomineral with greater porosity and a larger void space. For bivalves, CaCO3 crystal deposition is more chaotic in response to both ocean and coastal acidification. Biomechanics tests reveal that these changes result in weaker, more fragile skeletons, compromising their vital protective roles. Vulnerabilities differ among taxa and depend on acidification level. Climate warming has the potential to ameliorate some of the negative effects of acidification but may also make matters worse. The integrative morphology-ecomechanics approach is key to understanding how marine biominerals will perform in the face of changing climate

Ocean acidification impacts mussel control on biomineralisation

Ocean acidification is altering the oceanic carbonate saturation state and threatening the survival of marine calcifying organisms. Production of their calcium carbonate exoskeletons is dependent not only on the environmental seawater carbonate chemistry but also the ability to produce biominerals through proteins. We present shell growth and structural responses by the economically important marine calcifier Mytilus edulis to ocean acidification scenarios (380, 550, 750, 1000≈ atm pCO 2). After six months of incubation at 750≈ atm pCO 2, reduced carbonic anhydrase protein activity and shell growth occurs in M. edulis. Beyond that, at 1000≈ atm pCO 2, biomineralisation continued but with compensated metabolism of proteins and increased calcite growth. Mussel growth occurs at a cost to the structural integrity of the shell due to structural disorientation of calcite crystals. This loss of structural integrity could impact mussel shell strength and reduce protection from predators and changing environments

Crystallographic Interdigitation in Oyster Shell Folia Enhances Material Strength

Shells of oyster species belonging to the genus Crassostrea have similar shell microstructural features comprising well-ordered calcite folia. However, the mechanical strengths of folia differ dramatically between closely related species. For example, the calcareous shells of the Hong Kong oyster Crassostrea hongkongensis are stronger than those of its closest relative, the Portuguese oyster, Crassostrea angulata. Specifically, after removal of organic content, the folia of C. hongkongensis are 200% tougher and able to withstand a 100% higher crushing force than that of C. angulata. Detailed analyses of shell structural and mechanical features support the hypothesis that crystallographic interdigitations confer elevated mechanical strength in C. hongkongensis oyster shells compared to C. angulata shells. Consequently, the folia of C. hongkongensis are structurally equipped to withstand a higher external load compared to C. angulata. The observed relationships between oyster shell structure, crystallography, and mechanical properties provided an insightful context in which to consider the likely fate of these two species in future climate change scenarios. Furthermore, the interdisciplinary approach developed in this study through integrating electron backscatter diffraction (EBSD) data into finite element analysis (FEA) could be applied to other biomineral systems to investigate the relationship between crystallography and mechanical behavior

Increased pCO2 changes the lipid production in important aquacultural feedstock algae Isochrysis galbana, but not in Tetraselmis suecica

Increased anthropogenic CO2 emissions are leading to an increase in CO2 uptake by the world's oceans and seas, resulting in ocean acidification with a decrease in global ocean water pH by as much as 0.3–0.4 units by the year 2100. The direct effects of changing pCO2 on important microalgal feedstocks are not as well understood. Few studies have focused on lipid composition changes in specific algal species in response to ocean acidification and yet microalgae are an indispensable food source for various marine species, including juvenile shellfish. Isochrysis galbana and Tetraselmis suecica are widely used in aquaculture as feeds for mussels and other shellfish. The total lipid contents and concentrations of I. galbana and T. suecica were investigated when grown under present day (400 ppm) and ocean acidification conditions (1000 ppm) to elucidate the impact of increasing pCO2 on an important algae feedstock. Total lipids, long-chain alkenones (LCAs) and alkenoates decreased at 1000 ppm in I. galbana. I. galbana produces higher lipids than T. suecica, and is perhaps as a result more impacted by the change in carbon available for lipid production under higher pCO2. I. galbana is an important feedstock, more easily assimilated for growth in juvenile shellfish and reductions in lipid composition may prove problematic for the growth of future shellfish aquaculture. Our findings suggest that higher pCO2 impacts on algal lipid growth are species specific and warrant further study. It is therefore vital to examine the impact of high CO2 on algal lipid production, especially those commercial shellfish feed varieties to predict future impacts on commercial aquaculture

Biomineral shell formation under ocean acidification: A shift from order to chaos

Biomineral production in marine organisms employs transient phases of amorphous calcium carbonate (ACC) in the construction of crystalline shells. Increasing seawater pCO2 leads to ocean acidification (OA) with a reduction in oceanic carbonate concentration which could have a negative impact on shell formation and therefore survival. We demonstrate significant changes in the hydrated and dehydrated forms of ACC in the aragonite and calcite layers of Mytilus edulis shells cultured under acidification conditions (1000 μatm pCO2) compared to present day conditions (380 μatm pCO2). In OA conditions, Mytilus edulis has more ACC at crystalisation sites. Here, we use the high-spatial resolution of synchrotron X-ray Photo Emission Electron Microscopy (XPEEM) combined with X-ray Absorption Spectroscopy (XAS) to investigate the influence of OA on the ACC formation in the shells of adult Mytilus edulis. Electron Backscatter Diffraction (EBSD) confirms that OA reduces crystallographic control of shell formation. The results demonstrate that OA induces more ACC formation and less crystallographic control in mussels suggesting that ACC is used as a repair mechanism to combat shell damage under OA. However, the resultant reduced crystallographic control in mussels raises concerns for shell protective function under predation and changing environments. © 2016, Nature Publishing Group. All rights reserved

Selectively bred oysters can alter their biomineralization pathways, promoting resilience to environmental acidification

Commercial shellfish aquaculture is vulnerable to the impacts of ocean acidification driven by increasing carbon dioxide (CO2) absorption by the ocean as well as to coastal acidification driven by land run off and rising sea level. These drivers of environmental acidification have deleterious effects on biomineralization. We investigated shell biomineralization of selectively bred and wild‐type families of the Sydney rock oyster Saccostrea glomerata in a study of oysters being farmed in estuaries at aquaculture leases differing in environmental acidification. The contrasting estuarine pH regimes enabled us to determine the mechanisms of shell growth and the vulnerability of this species to contemporary environmental acidification. Determination of the source of carbon, the mechanism of carbon uptake and use of carbon in biomineral formation are key to understanding the vulnerability of shellfish aquaculture to contemporary and future environmental acidification. We, therefore, characterized the crystallography and carbon uptake in the shells of S. glomerata, resident in habitats subjected to coastal acidification, using high‐resolution electron backscatter diffraction and carbon isotope analyses (as δ13C). We show that oyster families selectively bred for fast growth and families selected for disease resistance can alter their mechanisms of calcite crystal biomineralization, promoting resilience to acidification. The responses of S. glomerata to acidification in their estuarine habitat provide key insights into mechanisms of mollusc shell growth under future climate change conditions. Importantly, we show that selective breeding in oysters is likely to be an important global mitigation strategy for sustainable shellfish aquaculture to withstand future climate‐driven change to habitat acidification

Chapter 2 Established and Emerging Techniques for Characterising the Formation, Structure and Performance of Calcified Structures under Ocean Acidification

Ocean acidification (OA) is the decline in seawater pH and saturation levels of calcium carbonate (CaCO3) minerals that has led to concerns for calcifying organisms such as corals, oysters and mussels because of the adverse effects of OA on their biomineralisation, shells and skeletons. A range of cellular biology, geochemistry and materials science approaches have been used to explore biomineralisation. These techniques have revealed that responses to seawater acidification can be highly variable among species, yet the underlying mechanisms remain largely unresolved. To assess the impacts of global OA, researchers will need to apply a range of tools developed across disciplines, many of which are emerging and have not yet been used in this context. This review outlines techniques that could be applied to study OA-induced alterations in the mechanisms of biomineralisation and their ultimate effects on shells and skeletons. We illustrate how to characterise, quantify and monitor the process of biomineralisation in the context of global climate change and OA. We highlight the basic principles, as well as the advantages and disadvantages, of established, emerging and future techniques for OA researchers. A combination of these techniques will enable a holistic approach and better understanding of the potential impact of OA on biomineralisation and its consequences for marine calcifiers and associated ecosystems

The response of coral skeletal nano-structure and hardness to ocean acidification conditions

Funding: Scottish Funding Council - HR09011; UK Natural Environment Research Council - NE/I022973/1.Ocean acidification typically reduces coral calcification rates and can fundamentally alter skeletal morphology. We use atomic force microscopy (AFM) and microindentation to determine how seawater pCO2 affects skeletal structure and Vickers hardness in a Porites lutea coral. At 400 µatm, the skeletal fasciculi are composed of tightly packed bundles of acicular crystals composed of quadrilateral nanograins, approximately 80–300 nm in dimensions. We interpret high adhesion at the nanograin edges as an organic coating. At 750 µatm the crystals are less regular in width and orientation and composed of either smaller/more rounded nanograins than observed at 400 µatm or of larger areas with little variation in adhesion. Coral aragonite may form via ion-by-ion attachment to the existing skeleton or via conversion of amorphous calcium carbonate precursors. Changes in nanoparticle morphology could reflect variations in the sizes of nanoparticles produced by each crystallization pathway or in the contributions of each pathway to biomineralization. We observe no significant variation in Vickers hardness between skeletons cultured at different seawater pCO2. Either the nanograin size does not affect skeletal hardness or the effect is offset by other changes in the skeleton, e.g. increases in skeletal organic material as reported in previous studies.Publisher PDFPeer reviewe

Ocean acidification reduces mechanical properties of the Portuguese oyster shell with impaired microstructure: a hierarchical analysis

Abstract. The rapidly intensifying process of ocean acidification (OA) in coastal areas due to anthropogenic CO2 is not only depleting carbonate ions necessary for calcification but also causing acidosis and disrupting internal pH homeostasis in several marine organisms. These negative consequences of OA on marine communities, particularly to shellfish oyster species, has been very well documented in recent studies, however, the consequences of these reduced or impaired calcification processes on the end-product, shells or skeletons, still remains one of the major research gaps. Shells produced by marine organisms under OA are expected to be corroded with disorganized or impaired crystal orientation or microstructures with reduced mechanical property. To bridge this knowledge gap and to test the above hypothesis, we investigated the effect of OA on shell of the commercially important oyster species (Crassostrea angulata) at ecologically and climatically relevant OA levels (using pH 8.1, 7.8, 7.5, 7.2 as proxies). In decreased pH conditions, a drop of shell hardness and stiffness was revealed by nanoindentation tests, while an evident loosened internal microstructure was detected by scanning electron microscopy (SEM). In contrary, the crystallographic orientation of oyster shell showed no significant difference with decreasing pH by Electron Back Scattered Diffraction (EBSD) analyses. These results indicate the loosened internal microstructure may be the cause of the OA induced reduction in shell hardness and stiffness. Micro-computed tomography analysis (Micro-CT) indicated that an overall "down-shifting" of mineral density in the shell with decreasing pH, which implied the loosened internal microstructure may run through the shell, thus inevitably limiting the effectiveness of the shell defensive function. This study surfaces potential bottom-up deterioration induced by OA on oyster shells, especially in their early juvenile life stage. This knowledge is critical to forecast the survival and production of edible oysters in future ocean. </jats:p