The requirement for calcification differs between ecologically important coccolithophore species.

Coccolithophores are globally distributed unicellular marine algae that are characterized by their covering of calcite coccoliths. Calcification by coccolithophores contributes significantly to global biogeochemical cycles. However, the physiological requirement for calcification remains poorly understood as non-calcifying strains of some commonly used model species, such as Emiliania huxleyi, grow normally in laboratory culture. To determine whether the requirement for calcification differs between coccolithophore species, we utilized multiple independent methodologies to disrupt calcification in two important species of coccolithophore: E. huxleyi and Coccolithus braarudii. We investigated their physiological response and used time-lapse imaging to visualize the processes of calcification and cell division in individual cells. Disruption of calcification resulted in major growth defects in C. braarudii, but not in E. huxleyi. We found no evidence that calcification supports photosynthesis in C. braarudii, but showed that an inability to maintain an intact coccosphere results in cell cycle arrest. We found that C. braarudii is very different from E. huxleyi as it exhibits an obligate requirement for calcification. The identification of a growth defect in C. braarudii resulting from disruption of the coccosphere may be important in considering their response to future changes in ocean carbonate chemistry.

Geographical CO2 sensitivity of phytoplankton correlates with ocean buffer capacity.

Accumulation of anthropogenic CO2 is significantly altering ocean chemistry. A range of biological impacts resulting from this oceanic CO2 accumulation are emerging, however, the mechanisms responsible for observed differential susceptibility between organisms and across environmental settings remain obscure. A primary consequence of increased oceanic CO2 uptake is a decrease in the carbonate system buffer capacity, which characterizes the system's chemical resilience to changes in CO2 , generating the potential for enhanced variability in pCO2 and the concentration of carbonate [ CO32- ], bicarbonate [ HCO3- ], and protons [H+ ] in the future ocean. We conducted a meta-analysis of 17 shipboard manipulation experiments performed across three distinct geographical regions that encompassed a wide range of environmental conditions from European temperate seas to Arctic and Southern oceans. These data demonstrated a correlation between the magnitude of natural phytoplankton community biological responses to short-term CO2 changes and variability in the local buffer capacity across ocean basin scales. Specifically, short-term suppression of small phytoplankton (<10 µm) net growth rates were consistently observed under enhanced pCO2 within experiments performed in regions with higher ambient buffer capacity. The results further highlight the relevance of phytoplankton cell size for the impacts of enhanced pCO2 in both the modern and future ocean. Specifically, cell size-related acclimation and adaptation to regional environmental variability, as characterized by buffer capacity, likely influences interactions between primary producers and carbonate chemistry over a range of spatio-temporal scales.

Why marine phytoplankton calcify.

Calcifying marine phytoplankton-coccolithophores- are some of the most successful yet enigmatic organisms in the ocean and are at risk from global change. To better understand how they will be affected, we need to know "why" coccolithophores calcify. We review coccolithophorid evolutionary history and cell biology as well as insights from recent experiments to provide a critical assessment of the costs and benefits of calcification. We conclude that calcification has high energy demands and that coccolithophores might have calcified initially to reduce grazing pressure but that additional benefits such as protection from photodamage and viral/bacterial attack further explain their high diversity and broad spectrum ecology. The cost-benefit aspect of these traits is illustrated by novel ecosystem modeling, although conclusive observations remain limited. In the future ocean, the trade-off between changing ecological and physiological costs of calcification and their benefits will ultimately decide how this important group is affected by ocean acidification and global warming.

Severity of ocean acidification following the end-Cretaceous asteroid impact.

Most paleo-episodes of ocean acidification (OA) were either too slow or too small to be instructive in predicting near-future impacts. The end-Cretaceous event (66 Mya) is intriguing in this regard, both because of its rapid onset and also because many pelagic calcifying species (including 100% of ammonites and more than 90% of calcareous nannoplankton and foraminifera) went extinct at this time. Here we evaluate whether extinction-level OA could feasibly have been produced by the asteroid impact. Carbon cycle box models were used to estimate OA consequences of (i) vaporization of up to 60 × 10(15) mol of sulfur from gypsum rocks at the point of impact; (ii) generation of up to 5 × 10(15) mol of NOx by the impact pressure wave and other sources; (iii) release of up to 6,500 Pg C as CO2 from vaporization of carbonate rocks, wildfires, and soil carbon decay; and (iv) ocean overturn bringing high-CO2 water to the surface. We find that the acidification produced by most processes is too weak to explain calcifier extinctions. Sulfuric acid additions could have made the surface ocean extremely undersaturated (Ωcalcite <0.5), but only if they reached the ocean very rapidly (over a few days) and if the quantity added was at the top end of literature estimates. We therefore conclude that severe ocean acidification might have been, but most likely was not, responsible for the great extinctions of planktonic calcifiers and ammonites at the end of the Cretaceous.

Predominance of heavily calcified coccolithophores at low CaCO3 saturation during winter in the Bay of Biscay.

Coccolithophores are an important component of the Earth system, and, as calcifiers, their possible susceptibility to ocean acidification is of major concern. Laboratory studies at enhanced pCO(2) levels have produced divergent results without overall consensus. However, it has been predicted from these studies that, although calcification may not be depressed in all species, acidification will produce "a transition in dominance from more to less heavily calcified coccolithophores" [Ridgwell A, et al., (2009) Biogeosciences 6:2611-2623]. A recent observational study [Beaufort L, et al., (2011) Nature 476:80-83] also suggested that coccolithophores are less calcified in more acidic conditions. We present the results of a large observational study of coccolithophore morphology in the Bay of Biscay. Samples were collected once a month for over a year, along a 1,000-km-long transect. Our data clearly show that there is a pronounced seasonality in the morphotypes of Emiliania huxleyi, the most abundant coccolithophore species. Whereas pH and CaCO(3) saturation are lowest in winter, the E. huxleyi population shifts from <10% (summer) to >90% (winter) of the heavily calcified form. However, it is unlikely that the shifts in carbonate chemistry alone caused the morphotype shift. Our finding that the most heavily calcified morphotype dominates when conditions are most acidic is contrary to the earlier predictions and raises further questions about the fate of coccolithophores in a high-CO(2) world.

Anthropogenic modification of the oceans.

Human activities are altering the ocean in many different ways. The surface ocean is warming and, as a result, it is becoming more stratified and sea level is rising. There is no clear evidence yet of a slowing in ocean circulation, although this is predicted for the future. As anthropogenic CO(2) permeates into the ocean, it is making sea water more acidic, to the detriment of surface corals and probably many other calcifiers. Once acidification reaches the deep ocean, it will become more corrosive to CaCO(3), leading to a considerable reduction in the amount of CaCO(3) accumulating on the deep seafloor. There will be a several thousand-year-long interruption to CaCO(3) sedimentation at many points on the seafloor. A curious feedback in the ocean, carbonate compensation, makes it more likely that global warming and sea-level rise will continue for many millennia after CO(2) emissions cease.

Eocene/Oligocene ocean de-acidification linked to Antarctic glaciation by sea-level fall.

One of the most dramatic perturbations to the Earth system during the past 100 million years was the rapid onset of Antarctic glaciation near the Eocene/Oligocene epoch boundary (approximately 34 million years ago). This climate transition was accompanied by a deepening of the calcite compensation depth--the ocean depth at which the rate of calcium carbonate input from surface waters equals the rate of dissolution. Changes in the global carbon cycle, rather than changes in continental configuration, have recently been proposed as the most likely root cause of Antarctic glaciation, but the mechanism linking glaciation to the deepening of calcite compensation depth remains unclear. Here we use a global biogeochemical box model to test competing hypotheses put forward to explain the Eocene/Oligocene transition. We find that, of the candidate hypotheses, only shelf to deep sea carbonate partitioning is capable of explaining the observed changes in both carbon isotope composition and calcium carbonate accumulation at the sea floor. In our simulations, glacioeustatic sea-level fall associated with the growth of Antarctic ice sheets permanently reduces global calcium carbonate accumulation on the continental shelves, leading to an increase in pelagic burial via permanent deepening of the calcite compensation depth. At the same time, fresh limestones are exposed to erosion, thus temporarily increasing global river inputs of dissolved carbonate and increasing seawater delta13C. Our work sheds new light on the mechanisms linking glaciation and ocean acidity change across arguably the most important climate transition of the Cenozoic era.

Phytoplankton calcification in a high-CO2 world.

Ocean acidification in response to rising atmospheric CO2 partial pressures is widely expected to reduce calcification by marine organisms. From the mid-Mesozoic, coccolithophores have been major calcium carbonate producers in the world's oceans, today accounting for about a third of the total marine CaCO3 production. Here, we present laboratory evidence that calcification and net primary production in the coccolithophore species Emiliania huxleyi are significantly increased by high CO2 partial pressures. Field evidence from the deep ocean is consistent with these laboratory conclusions, indicating that over the past 220 years there has been a 40% increase in average coccolith mass. Our findings show that coccolithophores are already responding and will probably continue to respond to rising atmospheric CO2 partial pressures, which has important implications for biogeochemical modeling of future oceans and climate.

Optical modeling and measurements of a coccolithophore bloom.

Blooms of the phytoplankton coccolithophorid Emiliania huxleyi can cause significant changes to both the inherent and the apparent optical properties within an oceanic column. Measurements made within such a bloom off the southwestern coast of England during July 1999 are reported. The multiple scattering properties of the bloom prevented accurate retrieval of absorption (a) and attenuation (c) coefficients with a WETLabs ac-9. Upwelling radiance measurements were similarly affected by the bloom, which caused the sensors to saturate. An optical model has been developed that gives close agreement with the in situ optics when it is used as input to the Hydrolight radiative-transfer model.