ALBERT

Description: Ocean acidification and warming are predicted to affect the ability of marine bivalves to build their shells, but little is known about the underlying mechanisms. Shell formation is an extremely complex process requiring a detailed understanding of biomineralization processes. Sodium incorporation into the shells would increase if bivalves rely on the exchange of Na+/H+ to maintain homeostasis for shell formation, thereby shedding new light on the acid-base and ionic regulation at the calcifying front. Here, we investigated the combined effects of seawater pH (8.1, 7.7 and 7.4) and temperature (16 and 22 °C) on the growth and sodium composition of the shells of the blue mussel, Mytilus edulis, and the Yesso scallop, Patinopecten yessoensis. Exposure of M. edulis to low pH (7.7 and 7.4) caused a significant decrease of shell formation, whereas a 6 °C warming significantly depressed the rate of shell growth in P. yessoensis. On the other hand, while the amount of Na incorporated into the shells of P. yessoensis did not increase in acidified seawater, an increase of Na/Cashell with decreasing pH was observed in M. edulis, the latter agreeing well with the aforementioned hypothesis. Moreover, a combined analysis of the shell growth and sodium content provides a more detailed understanding of shell formation processes. Under acidified conditions, mussels may maintain more alkaline conditions favorable for calcification, but a significant decrease of shell formation indicates that the mineralization processes are impaired. The opposite occurs in scallops; virtually unaffected shell growth implies that shell mineralization functions well. Finding of the present study may pave the way for deciphering the mechanisms underlying the impacts of ocean acidification and warming on bivalve shell formation.

Description: Ocean acidification is likely to have profound impacts on marine bivalves, especially on their early life stages. Therefore, it is imperative to know whether and to what extent bivalves will be able to acclimate or adapt to an acidifying ocean over multiple generations. Here, we show that reduced seawater pH projected for the end of this century (i.e., pH 7.7) led to a significant decrease of shell production of newly settled juvenile Manila clams, Ruditapes philippinarum. However, juveniles from parents exposed to low pH grew significantly faster than those from parents grown at ambient pH, exhibiting a rapid transgenerational acclimation to an acidic environment. The sodium composition of the shells may shed new light on the mechanisms responsible for beneficial transgenerational acclimation. Irrespective of parental exposure, the amount of Na incorporated into shells increased with decreasing pH, implying active removal of excessive protons through the Na+/H+ exchanger which is known to depend on the Na+ gradient actively built up by the Na+/K+-ATPase as a driving force. However, the shells with a prior history of transgenerational exposure to low pH recorded significantly lower amounts of Na than those with no history of acidic exposure. It therefore seems very likely that the clams may implement less costly and more ATP-efficient ion regulatory mechanisms to maintain pH homeostasis in the calcifying fluid following transgenerational acclimation. Our results suggest that marine bivalves may have a greater capacity to acclimate or adapt to ocean acidification by the end of this century than currently understood.

Description: Ocean acidification is predicted to have widespread implications for marine bivalve mollusks. While our understanding of its impact on their physiological and behavioral responses is increasing, little is known about their reproductive responses under future scenarios of anthropogenic climate change. In this study, we examined the physiological energetics of the Manila clam Ruditapes philippinarum exposed to CO2-induced seawater acidification during gonadal maturation. Three recirculating systems filled with 600 L of seawater were manipulated to three pH levels (8.0, 7.7, and 7.4) corresponding to control and projected pH levels for 2100 and 2300. In each system, temperature was gradually increased ca. 0.3 °C per day from 10 to 20 °C for 30 days and maintained at 20 °C for the following 40 days. Irrespective of seawater pH levels, clearance rate (CR), respiration rate (RR), ammonia excretion rate (ER), and scope for growth (SFG) increased after a 30-day stepwise warming protocol. When seawater pH was reduced, CR, ratio of oxygen to nitrogen, and SFG significantly decreased concurrently, whereas ammonia ER increased. RR was virtually unaffected under acidified conditions. Neither temperature nor acidification showed a significant effect on food absorption efficiency. Our findings indicate that energy is allocated away from reproduction under reduced seawater pH, potentially resulting in an impaired or suppressed reproductive function. This interpretation is based on the fact that spawning was induced in only 56% of the clams grown at pH 7.4. Seawater acidification can therefore potentially impair the physiological energetics and spawning capacity of R. philippinarum.

Description: Ocean acidification may interfere with the calcifying physiology of marine bivalves. Therefore, understanding their capacity for acclimation and adaption to low pH over multiple generations is crucial to make predictions about the fate of this economically and ecologically important fauna in an acidifying ocean. Transgenerational exposure to an acidification scenario projected by the end of the century (i.e., pH 7.7) has been shown to confer resilience to juvenile offspring of the Manila clam, Ruditapes philippinarum. However, whether, and to what extent, this resilience can persist into adulthood are unknown and the mechanisms driving transgenerational acclimation remain poorly understood. The present study takes observations of Manila clam juveniles further into the adult stage and observes similar transgenerational responses. Under acidified conditions, clams originating from parents reproductively exposed to the same level of low pH show a significantly faster shell growth rate, a higher condition index and a lower standard metabolic rate than those without prior history of transgenerational acclimation. Further analyses of stable carbon isotopic signatures in dissolved inorganic carbon of seawater, individual soft tissues and shells reveal that up to 61% of shell carbonate comes from metabolic carbon, suggesting that transgenerationally acclimated clams may preferentially extract internal metabolic carbon rather than transport external seawater inorganic carbon to build shells, the latter known to be energetically expensive. While a large metabolic carbon contribution (45%) is seen in non-acclimated clams, a significant reduction in the rate of shell growth indicates it might occur at the expense of other calcification-relevant processes. It therefore seems plausible that, following transgenerational acclimation, R. philippinarum can implement a less costly and more efficient energy-utilizing strategy to mitigate the impact of seawater acidification. Collectively, our findings indicate that marine bivalves are more resilient to ocean acidification projected for the end of the century than previously thought.