A review of transgenerational effects of ocean acidification on marine bivalves and their implications for sclerochronology

Ocean acidification can negatively impact marine bivalves, especially their shell mineralization processes. Consequently, whether marine bivalves can rapidly acclimate and eventually adapt in an acidifying ocean is now increasingly receiving considerable attention. Projecting the fate of this vulnerable taxonomic group is also pivotal for the science of sclerochronology - the study which seeks to deduce records of past environmental changes and organismal life-history traits from various geochemical properties of periodically layered hard tissues (bivalve shells, corals, fish otoliths, etc.). In this review, we provide a concise overview of the long-term and transgenerational responses of marine bivalves to elevated pCO(2) manifested at different levels of biological organization, with a specific focus on responses of geochemical properties (stable carbon and oxygen isotopes, minor and trace elements and microstructures) of their shells. Without exception, positive transgenerational responses to an elevated pCO(2) scenario projected for the year 2100 have been found in all five bivalve species hitherto studied, under the umbrella of two non-genetic mechanisms (increased maternal provisioning and epigenetic inheritance), suggesting that marine bivalves have remarkable transgenerational phenotypic plasticity which allows them to respond plastically and acclimate rapidly in an acidifying ocean. Rapid transgenerational acclimation, especially in terms of physiological processes, however, hinders a reliable interpretation of proxy records. Transgenerationally acclimated bivalves can actively modify the calcification physiology in response to elevated pCO(2), which in turn affects the processes of almost all geochemical proxies preserved in their shells. In particular, stable carbon isotopes, metabolically regulated elements (Na, K, Cu, Zn, Fe, etc.), and shell microstructures can be highly biased. In this context, we propose a number of challenges and opportunities the field of sclerochronology may face.