Decrease in coccolithophore calcification and CO2 since the middle Miocene.
ABSTRACT: Marine algae are instrumental in carbon cycling and atmospheric carbon dioxide (CO2) regulation. One group, coccolithophores, uses carbon to photosynthesize and to calcify, covering their cells with chalk platelets (coccoliths). How ocean acidification influences coccolithophore calcification is strongly debated, and the effects of carbonate chemistry changes in the geological past are poorly understood. This paper relates degree of coccolith calcification to cellular calcification, and presents the first records of size-normalized coccolith thickness spanning the last 14 Myr from tropical oceans. Degree of calcification was highest in the low-pH, high-CO2 Miocene ocean, but decreased significantly between 6 and 4 Myr ago. Based on this and concurrent trends in a new alkenone ?p record, we propose that decreasing CO2 partly drove the observed trend via reduced cellular bicarbonate allocation to calcification. This trend reversed in the late Pleistocene despite low CO2, suggesting an additional regulator of calcification such as alkalinity.
Project description:Calcite microfossils are widely used to study climate and oceanography in Earth's geological past. Coccoliths, readily preserved calcite plates produced by a group of single-celled surface-ocean dwelling algae called coccolithophores, have formed a significant fraction of marine sediments since the Late Triassic. However, unlike the shells of foraminifera, their zooplankton counterparts, coccoliths remain underused in palaeo-reconstructions. Precipitated in an intracellular chemical and isotopic microenvironment, coccolith calcite exhibits large and enigmatic departures from the isotopic composition of abiogenic calcite, known as vital effects. Here we show that the calcification to carbon fixation ratio determines whether coccolith calcite is isotopically heavier or lighter than abiogenic calcite, and that the size of the deviation is determined by the degree of carbon utilization. We discuss the theoretical potential for, and current limitations of, coccolith-based CO2 paleobarometry, that may eventually facilitate use of the ubiquitous and geologically extensive sedimentary archive.
Project description:Anthropogenic carbon dioxide emissions are forcing rapid ocean chemistry changes and causing ocean acidification (OA), which is of particular significance for calcifying organisms, including planktonic coccolithophores. Detailed analysis of coccolithophore skeletons enables comparison of calcite production in modern and fossil cells in order to investigate biomineralization response of ancient coccolithophores to climate change. Here we show that the two dominant coccolithophore taxa across the Paleocene-Eocene Thermal Maximum (PETM) OA global warming event (~56 million years ago) exhibited morphological response to environmental change and both showed reduced calcification rates. However, only Coccolithus pelagicus exhibits a transient thinning of coccoliths, immediately before the PETM, that may have been OA-induced. Changing coccolith thickness may affect calcite production more significantly in the dominant modern species Emiliania huxleyi, but, overall, these PETM records indicate that the environmental factors that govern taxonomic composition and growth rate will most strongly influence coccolithophore calcification response to anthropogenic change.
Project description:Background:The globally abundant coccolithophore, Emiliania huxleyi, plays an important ecological role in oceanic carbon biogeochemistry by forming a cellular covering of plate-like CaCO3 crystals (coccoliths) and fixing CO2. It is unknown how the cells arrange different-sized coccoliths to maintain full coverage, as the cell surface area of the cell changes during daily cycle. Methods:We used Euler's polyhedron formula and CaGe simulation software, validated with the geometries of coccoliths, to analyze and simulate the coccolith topology of the coccosphere and to explore the arrangement mechanisms. Results:There were only small variations in the geometries of coccoliths, even when the cells were cultured under variable light conditions. Because of geometric limits, small coccoliths tended to interlock with fewer and larger coccoliths, and vice versa. Consequently, to sustain a full coverage on the surface of cell, each coccolith was arranged to interlock with four to six others, which in turn led to each coccosphere contains at least six coccoliths. Conclusion:The number of coccoliths per coccosphere must keep pace with changes on the cell surface area as a result of photosynthesis, respiration and cell division. This study is an example of natural selection following Euler's polyhedral formula, in response to the challenge of maintaining a CaCO3 covering on coccolithophore cells as cell size changes.
Project description:Coccolithophorids are enigmatic plankton that produce calcium carbonate coccoliths, which over geological time have buried atmospheric CO2 into limestone, changing both the atmosphere and geology of the Earth. However, the role of coccoliths for the proliferation of these organisms remains unclear; suggestions include roles in anti-predation, enhanced photosynthesis and sun-screening. Here we test the hypothesis that calcification stabilizes the pH of the seawater proximate to the organisms, providing a level of acidification countering the detrimental basification that occurs during net photosynthesis. Such bioengineering provides a more stable pH environment for growth and fits the empirical evidence for changes in rates of calcification under different environmental conditions. Under this scenario, simulations suggest that the optimal production ratio of inorganic to organic particulate C (PIC : POCprod) will be lower (by approx. 20%) with ocean acidification and that overproduction of coccoliths in a future acidified ocean, where pH buffering is weaker, presents a risk to calcifying cells.
Project description:To understand the response of marine calcifying organisms under high CO2 scenarios, it is critical to study their calcification patterns in the natural environment. This paper focuses on a major calcifying phytoplankton group, the coccolithophores, through the analysis of water samples collected along a W-E Mediterranean transect during two research cruises, in April 2011 (Meteor cruise M84/3) and May 2013 (MedSeA cruise 2013). The Mediterranean Sea is a marginal sea characterized by large biogeochemical gradients. Currently, it is undergoing both warming and ocean acidification, processes which are rapidly modifying species distribution and calcification. The species Emiliania huxleyi largely dominates the total coccolithophore production in present day oceans and marine basins, including the Mediterranean Sea. A series of morphometric measurements were performed on the coccoliths of this species to estimate their mass, length and calculate a calcification index (proxy for the size-normalized calcification degree). The most abundant morphotype of E. huxleyi in the Mediterranean Sea is Type A. Coccoliths of this morphotype were additionally analyzed based on scanning electron microscopy images: four calcification varieties were quantified, according to the relationship between slit length-tube width, and the state of the central area (open or closed). The average E. huxleyi coccolith mass along the Mediterranean oceanographic transect depended strongly on both the average coccolith length and calcification index. The variability in average coccolith length and calcification index across samples reflected oscillations in the relative abundance of the calcification varieties. We also demonstrated that the distribution of the calcification varieties followed the main environmental gradients (carbonate chemistry, salinity, temperature, nutrient concentrations). Hence, shifts in the distribution of the calcification varieties and of the average E. huxleyi coccolith mass are to be expected in the Mediterranean Sea under climate change. These physiological and ecological responses will modulate the net coccolithophore calcification and, ultimately, the regional carbonate export to the seafloor.
Project description:Different morphotypes of the abundant marine calcifying algal species Emiliania huxleyi are commonly linked to various degrees of E. huxleyi calcification, but few studies have been done to validate this assumption. This study investigated therefore whether E. huxleyi morphotypes can be related to coccolithophore calcification and coccolith mass. Samples from January (high productivity) and September (low productivity) 1997 at an open ocean and a coastal site near the Canary Islands were analysed using a combination of thickness measurements (Circular Polarizer Retardation estimates (CPR) method), Scanning Electron Microscope imaging, and Markov Chain Monte Carlo (MCMC) models. Mean E. huxleyi coccolith mass varied from a maximum of 2.9pg at the open ocean station in January to a minimum of 1.7pg in September at both stations. In contrast, overall calcite produced by E. huxleyi (assuming 23 coccoliths/cell) varied from a maximum of 2.6 ?gL-1 at the coastal station in January to a minimum of 0.5 ?gL-1 in September at the open ocean site. The relative abundance of "Overcalcified" Type A, Type A, Group B and malformed coccoliths was determined from SEM images. The mean coccolith mass of "Overcalcified" Type A was 2.0pg using the CPR-method, while mean mass of Type A and Group B coccoliths was determined using coccolith length measurements from SEM images and MCMC models relating thickness measurements to morphotype relative abundance. Type A cocccolith mass varied from a 1.6pg to 2.6pg and Group B coccolith mass varied from 1.5pg to 2.0pg. These results demonstrate that the coccolith mass of Type A, "Overcalcified" Type A, and Group B do not differ systematically and there is no systematic relationship between relative abundance of a morphotype and the overall calcite production of E. huxleyi. Therefore, morphotype appearance and relative abundance can not be uniformly used as reliable indicators of E. huxleyi calcification or calcite production.
Project description:Coccolithophores are phytoplanktonic algae which produce an exoskeleton made of single platelets of calcite named coccoliths. They are widespread in all oceans and directly impact the short- and long-term C cycle. The study of coccolith size, morphology and elemental composition reveals important information regarding the ability of the cell to calcify and on the factors that influence this process. In this regard, very little is known about coccolith composition and its changes under altered environmental conditions. Here, we present high resolution (50 × 50?nm) elemental spatial distribution in pristine coccoliths of Coccolithus pelagicus and Gephyrocapsa oceanica reconstructed via X-ray fluorescence analyses at synchrotron. The studied specimens are from control culture and metal-enriched (V, Ni, Zn and Pb) experiments. The analysed specimens produced under stress conditions, display an irregular shape and are thinner, especially in the external rim, with ca. 1/3 lower Ca concentrations compared to specimens from the control. The same specimens also have higher Sr/Ca ratio with highest values in the coccolith external rim, suggesting that difficulty in calcification is additionally reflected in increased Sr/Ca ratios. Selenium is found in the coccolith as possible substitute of carbonate in the calcite. V and Pb apparently did not interact with the coccoliths while Zn and Ni were deposited on the coccolith surface.
Project description:Biomineralization by calcifying microalgae is a precisely controlled intracellular calcification process that produces delicate calcite scales (or coccoliths) in the coccolithophore Emiliania huxleyi (Haptophycea). Despite its importance in biogeochemical cycles and the marine environment globally, the underlying molecular mechanism of intracellular coccolith formation, which requires calcium, bicarbonate, and coccolith-polysaccharides, remains unclear. In E. huxleyi CCMP 371, we demonstrated that reducing the calcium concentration from 10 (ambient seawater) to 0.1 mM strongly restricted coccolith production, which was then recovered by adding 10 mM calcium, irrespective of inorganic phosphate conditions, indicating that coccolith production could be finely controlled by the calcium supply. Using this strain, we investigated the expression of differentially expressed genes (DEGs) to observe the cellular events induced by changes in calcium concentrations. Intriguingly, DEG analysis revealed that the phosphatidylinositol-specific phospholipase C (PI-PLC) gene was upregulated and coccolith production by cells was blocked by the PI-PLC inhibitor U73122 under conditions closely associated with calcium-induced calcification. These findings imply that PI-PLC plays an important role in the biomineralization process of the coccolithophore E. huxleyi.
Project description:Ocean acidification will potentially inhibit calcification by marine organisms; however, the response of the most prolific ocean calcifiers, coccolithophores, to this perturbation remains under characterized. Here we report novel chemical constraints on the response of the widespread coccolithophore species Ochrosphaera neapolitana (O. neapolitana) to changing-CO2 conditions. We cultured this algae under three pCO2-controlled seawater pH conditions (8.05, 8.22, and 8.33). Boron isotopes within the algae's extracellular calcite plates show that this species maintains a constant pH at the calcification site, regardless of CO2-induced changes in pH of the surrounding seawater. Carbon and oxygen isotopes in the algae's calcite plates and carbon isotopes in the algae's organic matter suggest that O. neapolitana utilize carbon from a single internal dissolved inorganic carbon (DIC) pool for both calcification and photosynthesis, and that a greater proportion of dissolved CO2 relative to HCO3- enters the internal DIC pool under acidified conditions. These two observations may explain how O. neapolitana continues calcifying and photosynthesizing at a constant rate under different atmospheric-pCO2 conditions.
Project description:In this study, we present experimental evidence showing that coccoliths have light-scattering anisotropy that contributes to a possible control of solar light exposure in the ocean. Changing the angle between the incident light and an applied magnetic field causes differences in the light-scattering intensities of a suspension of coccoliths isolated from Emiliania huxleyi. The magnetic field effect is induced by the diamagnetic torque force directing the coccolith radial plane perpendicular to the applied magnetic fields at 400 to 500 mT. The developed technique reveals the light-scattering anisotropies in the 3-μm-diameter floating coccoliths by orienting themselves in response to the magnetic fields. The detached coccolith scatters radially the light incident to its radial plane. The experimental results on magnetically oriented coccoliths show that an individual coccolith has a specific direction of light scattering, although the possible physiological effect of the coccolith remains for further study, focusing on the light-scattering anisotropies of coccoliths on living cells.