carbon cycle. Coccolithophores are the major planktonic group responsible for pelagic CaCO3 production. The diagram on the right shows the energetic costs of coccolithophore calcification: (A) Transport processes include the transport into the cell from the surrounding seawater of primary calcification substrates Ca2+ and HCO3− (black arrows) and the removal of the end product H+ from the cell (gray arrow). The transport of Ca2+ through the cytoplasm to the CV is the dominant cost associated with calcification. (B) Metabolic processes include the synthesis of CAPs (gray rectangles) by the Golgi complex (white rectangles) that regulate the nucleation and geometry of CaCO3 crystals. The completed coccolith (gray plate) is a complex structure of intricately arranged CAPs and CaCO3 crystals. (C) Mechanical and structural processes account for the secretion of the completed coccoliths that are transported from their original position adjacent to the nucleus to the cell periphery, where they are transferred to the surface of the cell. The costs associated with these processes are likely to be comparable to organic-scale exocytosis in noncalcifying haptophyte algae. The diagram on the left shows the benefits of coccolithophore calcification. (A) Accelerated photosynthesis includes CCM (1) and enhanced light uptake via scattering of scarce photons for deep-dwelling species (2). (B) Protection from photodamage includes sunshade protection from ultraviolet (UV) light and photosynthetic active radiation (PAR) (1) and energy dissipation under high-light conditions (2). (C) Armor protection includes protection against viral/bacterial infections (1) and grazing by selective (2) and nonselective (3) grazers. The degree by which calcification can adapt to ocean acidification is presently unknown. Cell physiological examinations found the essential H+ efflux (stemming from the use of HCO3− for intra-cellular calcification) to become more costly with ongoing ocean acidification as the electrochemical H+ inside-out gradient is reduced and passive proton outflow impeded. Adapted cells would have to activate proton channels more frequently, adjust their membrane potential, and/or lower their internal pH. Reduced intra-cellular pH would severely affect the entire cellular machinery and require other processes (e.g. photosynthesis) to co-adapt in order to keep H+ efflux alive. The obligatory H+ efflux associated with calcification may therefore pose a fundamental constraint on adaptation which may potentially explain why "calcification crisis" were possible during long-lasting (thousands of years) CO2 perturbation events even though evolutionary adaption to changing carbonate chemistry conditions is possible within one year. Unraveling these fundamental constraints and the limits of adaptation should be a focus