Coccolith Production Research Articles

Calcification increases carbon supply, photosynthesis, and growth in a globally distributed coccolithophore

AbstractCoccolithophores fix organic carbon and produce calcite plates (coccoliths) that ballast organic matter and facilitate carbon export. Photosynthesis consumes carbon dioxide, while calcification produces it, raising questions about whether coccolithophores are a net sink or source of carbon. We characterized the physiology of calcified and noncalcified (“naked”) phenotypes of Emiliania huxleyi (CCMP374) and investigated the relationship between calcification and photosynthesis across a gradient of light (25–2000 μmol photons m−2 s−1) spanning the euphotic zone. Growth and photophysiological parameters increased with light until reaching a mid‐light (150 μmol photons m−2 s−1) maximum for both phenotypes. Calcified cells were characterized by enhanced photophysiology and less photoinhibition. Further, enhanced bicarbonate transport in calcified cells led to higher rates of particulate organic carbon fixation and growth compared to naked cells at mid‐light to high light (150–2000 μmol photons m−2 s−1). Coccolith production was similarly high at mid and high light, but the rate of coccolith shedding was >3‐fold lower at high‐light (1.2 vs. 0.35 coccoliths cell−1 h−1). The cellular mechanims(s) of this differential shedding remain unknown and underly light‐related controls on coccosphere maintenance. Our data suggest coccoliths shade cells at high light and that enhanced bicarbonate transport associated with calcification increases internal carbon supplies available for organic carbon fixation.

Variations in the Southern Ocean carbonate production, preservation, and hydrography for the past 41, 500 years: Evidence from coccolith and CaCO3 records

Changes in ocean alkalinity affect atmospheric pCO2 (i.e., higher alkalinity lowers atmospheric pCO2). Ocean alkalinity is partly determined by sedimentary burial of carbonates, which is primarily controlled by carbonate flux and the degree of deep ocean carbonate saturation. In this study, we investigate the factors determining the coccolith burial in subantarctic sediments and the surface ocean changes in the subtropical South Indian Ocean. The downcore coccolith records from the subantarctic region (SK200/22a) of the Indian sector of the Southern Ocean display low coccolith concentration during the glacial period. A possible explanation for this is, 1) the low glacial production of coccolithophores due to the competition from diatoms and 2) dilution by biogenic silica in the glacial sediments. Additionally, reduced carbonate burial owing to the low carbonate saturation of the deep-water accounts for the decline in glacial coccolith concentration. This also explains the low coccolith dissolution index and enrichment of the large dissolution-resistant coccolith species, Coccolithus pelagicus subsp. braarudii in the glacial sediments. The low carbonate saturation is attributed to, 1) the replacement of carbonate saturated, North Atlantic Deep Waters by the undersaturated southern sourced water masses and 2) increased storage of dissolved CO2 in the deep glacial Southern Ocean. Our study suggests that changes in coccolith production and the deep ocean carbonate saturation determine their burial in subantarctic sediments for the last 41,500 years. Other than these changes, the study region also records the changes in the Agulhas Return Current via variation in the proportion of tropical-subtropical coccolith assemblage.

Enhanced Carbonate Counter Pump and upwelling strengths in the Indian sector of the Southern Ocean during MIS 11

While numerous studies have highlighted the central role of Southern Ocean (SO) dynamics in modulating rapid increases in atmospheric CO2 concentrations during deglaciations, fewer studies have yet focused on the impact of the Biological Carbon Pump - and more specifically the Carbonate Counter Pump (CCP) - in contributing to increase the CO2 concentration in oceanic surface waters and thus, in the atmosphere. Here, we present micropaleontological (coccolith, planktonic foraminifera) and geochemical (CaCO3, CaXRF, δ13CN. pachyderma) constraints from sediment core MD04-2718 retrieved in the Polar Front Zone of the Indian Ocean covering the time interval spanning Marine Isotope Stage (MIS) 12 to MIS 10 (440,000–360,000 years). We compare our results with published records from the SO to reconstruct past changes in CCP and upwelling dynamics and understand their leverage on the ocean-atmosphere portioning of CO2. We demonstrate that the sharp increase in atmospheric pCO2 during Termination V was likely associated with enhanced deep-water ventilation in the SO, that promoted the release of previously sequestered CO2 to the ocean surface as the westerly wind belt and the frontal system migrated southwards. Enhanced CCP is observed later, during MIS 11, and is likely the consequence of higher sea surface temperature and higher nutrient availability due to the reinvigoration of SO upwelling leading to increased coccolith (and to a lesser degree, planktonic foraminifera) production and export. The low eccentricity signal recorded during MIS 11 might have additionally strengthened the CCP, exerting a specific control on Gephyrocapsa morphotypes. In addition to the strong global biological productivity and higher carbon storage on land, these synergistic mechanisms may have permitted to shape the distinctive 30 ka-long pCO2 plateau characteristic of MIS 11.

Coccolith size rules – What controls the size of coccoliths during coccolithogenesis?

Heterococcoliths are calcite platelets produced inside diploid coccolithophore cells and extruded to form a covering on the cell surface called a coccosphere. The size of coccoliths is an important parameter sometimes used to identify species, and it is observed to be influenced in extant species by abiotic parameters (e.g., CO2, light). However, the variable distribution of coccolith sizes occurring within a single coccosphere questions the mechanisms controlling coccolith size. A relationship between cell/coccosphere size and mean coccolith size was previously identified, called the “coccolithophore size rules”. In this study, we query the mechanisms controlling the size of a coccolith during coccolithogenesis. A culture experiment on Gephyrocapsa huxleyi strain RCC1216 shows that coccolithogenesis occurs during the cell growth G1 interphase and newly produced coccoliths get bigger as the cell grows. These observations provide parameters for the development of two numerical models used to simulate the coccolith size distribution within a coccolithophore population. Neither model can accurately reproduce an empirical monoclonal coccolith size distribution, indicating that additional factors influence coccolith size. According to our results, coccolith size is only clearly related to cell size at the time of its formation. We confirm that application of the coccolithophore size rules model should be limited to inferring average cell dimensions from (fossil) coccolith biometry, and that comparisons are valid only in multipopulational studies. The coccolith size rule model – the constraining effect of coccolith production during G1 interphase cell growth on coccolith size – proposed here is applicable only for some placolith-forming species.

Association of Phosphatidylinositol-Specific Phospholipase C with Calcium-Induced Biomineralization in the Coccolithophore Emiliania huxleyi.

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.

Sr in coccoliths of Scyphosphaera apsteinii: Partitioning behavior and role in coccolith morphogenesis

Coccolithophores are important contributors to global calcium carbonate through their species-specific production of calcite coccoliths. Nannofossil coccolith calcite remains an important tool for paleoreconstructions through geochemical analysis of isotopic and trace element incorporation including Sr, which is a potential indicator of past surface ocean temperature and productivity. Scyphosphaera apsteinii (Zygodiscales) exhibits an unusually high Sr/Ca ratio and correspondingly high partitioning coefficient (DSr = 2.5) in their two morphologically distinct types of coccoliths: flat muroliths and barrel-like lopadoliths. Whether or not this reflects mechanistic differences in calcification compared to other coccolithophores is unknown. We therefore examined the possible role of Sr in S. apsteinii calcification by growing cells in deplete (0.33 mmol/mol Sr/Ca), ambient (9 mmol/mol Sr/Ca), and higher than ambient Sr conditions (36 and 72 mmol/mol Sr/Ca). The effects on growth, quantum efficiency of photosystem II (Fv/Fm), coccolith morphology, and calcite DSr were evaluated. No effect on S. apsteinii growth rate or Fv/Fm was observed when cells were grown in Sr/Ca between 0.33–36 mmol/mol. However, at 72 mmol/mol Sr/Ca growth rate was significantly reduced, although Fv/Fm was unaffected. Reducing the Sr/Ca from ambient (9 mmol/mol) did not significantly alter the frequency of malformed and aberrant muroliths and lopadoliths, but at higher than ambient Sr/Ca conditions coccolith morphology was significantly disrupted. This implies that Sr is not a critical determining factor in normal coccolith calcite morphology in this dimorphic species. Using energy dispersive spectroscopy (EDS) we observed an increase in [Sr] and decrease in DSr of coccoliths as the Sr/Ca of the growth medium increased. Interestingly, muroliths had significantly lower Sr/Ca than lopadoliths at ambient and elevated [Sr], and lopadolith tips had lower Sr than bases in ambient conditions. In summary, the Sr fractionation behavior of S. apsteinii is distinct from other coccolithophores because of an unusually high DSr and inter- and intra-coccolith variability in Sr/Ca. These observations could be explained by mechanistic differences in the selectivity of the Ca2+ transport pathway or in the Sr-and Ca-binding capacity of organic components, such as polysaccharides associated with coccolithogenesis.

Reduced continental weathering and marine calcification linked to late Neogene decline in atmospheric CO2

The globally averaged calcite compensation depth has deepened by several hundred metres in the past 15 Myr. This deepening has previously been interpreted to reflect increased alkalinity supply to the ocean driven by enhanced continental weathering due to the Himalayan orogeny during the late Neogene period. Here we examine mass accumulation rates of the main marine calcifying groups and show that global accumulation of pelagic carbonates has decreased from the late Miocene epoch to the late Pleistocene epoch even though CaCO3 preservation has improved, suggesting a decrease in weathering alkalinity input to the ocean, thus opposing expectations from the Himalayan uplift hypothesis. Instead, changes in relative contributions of coccoliths and planktonic foraminifera to the pelagic carbonates in relative shallow sites, where dissolution has not taken its toll, suggest that coccolith production in the euphotic zone decreased concomitantly with the reduction in weathering alkalinity inputs as registered by the decline in pelagic carbonate accumulation. Our work highlights a mechanism whereby, in addition to deep-sea dissolution, changes in marine calcification acted to modulate carbonate compensation in response to reduced weathering linked to the late Neogene cooling and decline in atmospheric partial pressure of carbon dioxide. A redistribution of marine calcifiers along with a reduction in weathering led to increased seafloor carbonate deposition during the late Neogene, according to a global compilation of carbonate mass accumulation rate records from sediment cores.

Deciphering latitudinal shifts in coccolith accumulation in the eastern tropical Pacific Ocean through the Pleistocene

Abstract Middle and Late Pleistocene coccolithophore assemblages from Ocean Drilling Program (ODP) sites 1241 and 1242 in the northeastern tropical Pacific (north ETP) were analyzed to reconstruct coccolithophore-productivity and surface-water conditions over the last 925 kyr. Stratigraphic control was provided by δ18O data and geomagnetic time scales and the identification of synchronous calcareous nannofossil events, formerly described in mid and low latitudes. Changes in sea surface coccolithophore-productivity were inferred by estimates of the so-called N ratio – an independent proxy on conversion to accumulation rates – and the total coccolith-derived carbonate accumulation rate (CAR). CAR variations follow those of the N ratio but show larger amplitude changes. Fluctuations in the N ratio and total CAR values imply coccolithophore-productivity maxima during glacial times, most likely as a result of a shallower nutri-thermocline, defined as the boundary beneath the sea surface at which nutrient-rich/cool and nutrient-poor/warm waters are separated. Low interglacial values of the N ratio and CAR suggest lower coccolithophore-productivity and a deeper nutri-thermocline (deep stratification of the mixed layer). This result implies that coccolithophore-productivity-controlled CARs and dissolution were driven by the ratio between CaCO3 production and organic matter oxidation, and that effects of dissolution were enhanced when coccolith production fell. The data suggest high coccolithophore-productivity and a shallow nutri-thermocline during MIS 22 through MIS 6, likely due to the reinforcement of northeastern wind-driven advection of nutrient-rich waters, which is compatible with a La Nina-like state. The dominance of Gephyrocapsa caribbeanica during the Mid-Brunhes Event (MBE) is consistent with this eutrophic environment. Conversely, data from MIS 5 suggests a weakening of trade winds in the ETP, which would have reduced the normal upwelling of cool water and warmed SST, and which is compatible with the oligotrophic setting of an El Nino-like state. During MIS 4 through MIS 1, coccolithophore-productivity was not significantly different from MIS 5 and therefore not indicative of a prominent La Nina-like state, as previously reported for the Equatorial Cold Tongue (ECT). Comparison with coccolithophore data from Site 1240 supports a reduction of coccolithophore-productivity and nutri-thermocline deepening from the ECT to the Costa Rica margin and suggests that nutrient availability was the primary control on the distribution of the whole assemblage during the past 925 kyr.

Paleoceanographic evolution of the Japan Sea over the last 460 kyr – A coccolithophore perspective

Abstract Changes in the intensity of the influx of the Tsushima Warm Current (TWC) in the south-central part of the Japan Sea (JS), or East Sea in Korean, were reconstructed for the last 460 kyr at IODP Site U1427 using the composition and abundance of the coccolithophore assemblage. In addition, the recent distribution of coccolithophore taxa in the JS and the East China Sea was assessed using electron microscopy. Coccolithophore assemblages, dominated by Emiliania huxleyi and Gephyrocapsa species, and coccolith abundance reveal a strong relationship between sea level and ocean variability over the last five glacial/interglacial cycles, in good agreement with planktic foraminiferal data. Three different circulation modes based on calcareous nannofossil and foraminifera TWC indicators were proposed for the JS. Coccolith production was low and TWC indicators (i.e., Gephyrocapsa oceanica, Calcidiscus leptoporus s.l., and Helicosphaera carteri) absent due to the isolation of the JS during glacials in response to global sea level falls (Mode 1). In contrast, coccolith abundance and TWC indicators reach a maximum due to the most intense TWC flow through the Tsushima Strait during interglacials (Mode 3). Intermediate conditions (Mode 2) are characterized by moderate/high coccolith numbers, presence of TWC coccolith indicators, and rare TWC planktic foraminifera indicators. This mode resulted in intermittent variations in the contribution of the TWC and East China Sea coastal water due to relatively low sea level stands (ca. −90 m to −20 m). Coccoliths dominated the carbonate sequence prior to MIS 8 and were major contributors to the total carbonate of the sediment at Site U1427, suggesting high coccolithophore productivity and a bloom-type environment during MIS 11 and 9. These changes in the carbonate chemistry caused by glacio-eustacy in the JS at the northwest margin of the Pacific Ocean should be considered in future paleoceanographic and paleoclimate models.

Formation and Separation of Particles from Emiliania huxleyi

AbstractThree reactor systems for Emiliania huxleyi cultivation are reviewed in respect to coccolith production. After cell lysis, the particle size distribution was determined for the system. Fraction changes caused through ultrasonication were characterized by comparing particle size distributions. Insufficient de‐agglomeration and possible re‐agglomeration at longer treatment intervals could be observed. Zeta potential measurements of the treated samples indicated no significant surface modifications. Scanning electron microscopy images showed no structural damages and the particles appeared similar to each other regardless the ultrasound exposure. In addition, settling velocities of the system using different density and viscosity solutions were examined by an analytical centrifuge. Two separation attempts for coccolith recovery are presented.

Growth and mortality of coccolithophores during spring in a temperate Shelf Sea (Celtic Sea, April 2015)

Coccolithophores are key components of phytoplankton communities, exerting a critical impact on the global carbon cycle and the Earth’s climate through the production of coccoliths made of calcium carbonate (calcite) and bioactive gases. Microzooplankton grazing is an important mortality factor in coccolithophore blooms, however little is currently known regarding the mortality (or growth) rates within non-bloom populations. Measurements of coccolithophore calcite production (CP) and dilution experiments to determine microzooplankton (≤63 µm) grazing rates were made during a spring cruise (April 2015) at the Central Celtic Sea (CCS), shelf edge (CS2), and within an adjacent April bloom of the coccolithophore Emiliania huxleyi at station J2.CP at CCS ranged from 10.4 to 40.4 µmol C m−3 d−1 and peaked at the height of the spring phytoplankton bloom (peak chlorophyll-a concentrations ∼6 mg m−3). Cell normalised calcification rates declined from ∼1.7 to ∼0.2 pmol C cell−1 d−1, accompanied by a shift from a mixed coccolithophore species community to one dominated by the more lightly calcified species E. huxleyi and Calciopappus caudatus. At the CCS, coccolithophore abundance increased from 6 to 94 cells mL−1, with net growth rates ranging from 0.06 to 0.21 d−1 from the 4th to the 28th April. Estimates of intrinsic growth and grazing rates from dilution experiments, at the CCS ranged from 0.01 to 0.86 d−1 and from 0.01 to 1.32 d−1, respectively, which resulted in variable net growth rates during April. Microzooplankton grazers consumed 59 to >100% of daily calcite production at the CCS. Within the E. huxleyi bloom a maximum density of 1986 cells mL−1 was recorded, along with CP rates of 6000 µmol C m−3 d−1 and an intrinsic growth rate of 0.29 d−1, with ∼80% of daily calcite production being consumed.Our results show that microzooplankton can exert strong top-down control on both bloom and non-bloom coccolithophore populations, grazing over 60% of daily growth (and calcite production). The fate of consumed calcite is unclear, but may be lost either through dissolution in acidic food vacuoles, and subsequent release as CO2, or export to the seabed after incorporation into small faecal pellets. With such high microzooplankton-mediated mortality losses, the fate of grazed calcite is clearly a high priority research direction.

A Conceptual Model for Projecting Coccolithophorid Growth, Calcification and Photosynthetic Carbon Fixation Rates in Response to Global Ocean Change

Temperature, light and carbonate chemistry speciation all influence the growth, calcification and photosynthetic carbon fixation rates of coccolithophores to a similar degree. There have been multiple attempts to project the responses of coccolithophores to changes in carbonate chemistry, but the interaction with light and temperature remains elusive. Here we devise a simple conceptual model to derive a fit equation for coccolithophorid growth, photosynthetic carbon fixation and calcification rates in response to simultaneous changes in carbonate chemistry speciation, temperature and light conditions. The fit equation is able to account for up to 88% of the variability in measured metabolic rates. Equation projections indicate that temperature, light and carbonate chemistry speciation all have different modulating effects on both optimal growth conditions and the sensitivity of responses to extreme environmental conditions. Calculations suggest that a single extreme environmental condition (CO2, temperature, light) will reduce maximum rates regardless of how optimal the other environmental conditions may be. Thus, while the response of coccolithophores to ocean change depends on multiple variables, the one which is least optimal will have the most impact on overall rates. Finally, responses to ocean change are usually reported in terms of cellular rates. However, changes in cellular rates can be a poor predictor for assessing changes in production at the community level. We therefore introduce a new metric, the calcium carbonate production potential (CCPP), which combines the independent effects of changes in growth rate and cellular calcium carbonate content to assess how environmental changes will impact coccolith production. Direct comparison of CO2 impacts on cellular CaCO3 production rates and CCPP shows that while the former is still at 45% of its pre-industrial capacity at 1000 uatm, the latter is reduced to 10%.

Erratum to: Patterns of coccolithophore pigment change under global acidification conditions based on in-situ observations at BATS site between July 1990–Dec 2008

Coccolith production is an important part of the biogenic carbon cycle as the largest source of calcium carbonate on earth, accounting for about 75% of the deposition of carbon on the sea floor. Recent studies based on laboratory experiment results indicated that increasing anthropogenic CO2 in the atmosphere triggered global ocean acidification leading to a decrease of calcite or aragonite saturation and calcium carbonate, and to decreasing efficiency of carbon export/pumping to deep layers. In the present study, we analyzed about 20 years of field observations of coccolithophore pigment, dissolved inorganic carbon (DIC), nutrients, and temperatures from the Bermuda Atlantic Time-series Study (BATS) site and satellite remote sensing to investigate the variable tendency of the coccolithophore pigment, and to evaluate the influence of ocean acidification on coccolithophore biomass. The results indicated that there was a generally increasing tendency of coccolithophore pigment, coupled with increasing bicarbonate concentrations or decreasing carbonate ion concentration. The change of coccolithophore pigment was also closely associated with pH, nutrients, mixed layer depth (MLD), and temperature. Correlation analyses between coccolithophores and abiotic parameter imply that coccoliths production or coccolithophore pigment has increased with increasing acidification in the recent 20 years.

Environmental drivers of coccolithophore abundance and calcification across Drake Passage (Southern Ocean)

Abstract. Although coccolithophores are not as numerically common or as diverse in the Southern Ocean as they are in subpolar waters of the North Atlantic, a few species, such as Emiliania huxleyi, are found during the summer months. Little is actually known about the calcite production (CP) of these communities or how their distribution and physiology relate to environmental variables in this region. In February 2009, we made observations across Drake Passage (between South America and the Antarctic Peninsula) of coccolithophore distribution, CP, primary production, chlorophyll a and macronutrient concentrations, irradiance and carbonate chemistry. Although CP represented less than 1 % of total carbon fixation, coccolithophores were widespread across Drake Passage. The B/C morphotype of E. huxleyi was the dominant coccolithophore, with low estimates of coccolith calcite (∼ 0.01 pmol C coccolith−1) from biometric measurements. Both cell-normalised calcification (0.01–0.16 pmol C cell−1 d−1) and total CP (< 20 µmol C m−3 d−1) were much lower than those observed in the subpolar North Atlantic where E. huxleyi morphotype A is dominant. However, estimates of coccolith production rates were similar (0.1–1.2 coccoliths cell−1 h−1) to previous measurements made in the subpolar North Atlantic. A multivariate statistical approach found that temperature and irradiance together were best able to explain the observed variation in species distribution and abundance (Spearman's rank correlation ρ = 0.4, p < 0.01). Rates of calcification per cell and coccolith production, as well as community CP and E. huxleyi abundance, were all positively correlated (p < 0.05) to the strong latitudinal gradient in temperature, irradiance and calcite saturation states across Drake Passage. Broadly, our results lend support to recent suggestions that coccolithophores, especially E. huxleyi, are advancing polewards. However, our in situ observations indicate that this may owe more to sea-surface warming and increasing irradiance rather than increasing CO2 concentrations.

Coccolithophore Cell Biology: Chalking Up Progress.

Coccolithophores occupy a special position within the marine phytoplankton because of their production of intricate calcite scales, or coccoliths. Coccolithophores are major contributors to global ocean calcification and long-term carbon fluxes. The intracellular production of coccoliths requires modifications to cellular ultrastructure and metabolism that are surveyed here. In addition to calcification, which appears to have evolved with a diverse range of functions, several other remarkable features that likely underpin the ecological and evolutionary success of coccolithophores have recently been uncovered. These include complex and varied life cycle strategies related to abiotic and biotic interactions as well as a range of novel metabolic pathways and nutritional strategies. Together with knowledge of coccolithophore genetic and physiological variability, these findings are beginning to shed new light on species diversity, distribution, and ecological adaptation. Further advances in genetics and functional characterization at the cellular level will likely to lead to a rapid increase in this understanding.

Palaeoenvironmental vs. evolutionary control on size variation of coccoliths across the Lower-Middle Jurassic

Size is an easily quantifiable trait probing the respective influence of evolution and palaeoenvironment on organisms through time. After the stressed period that spanned across western Tethys epicontinental seas during the early Toarcian, the following rebuilding of a restored water column ecosystem is coeval with the size increase of the Lotharingius genus coccoliths, one of the main pelagic producers at that time. In order to quantitatively assess the coccolith size evolution during a time span of ~10myr, measurements of seven morphospecies of Lotharingius across the Toarcian and early Aalenian were performed. This study was carried out on a total of 5500 specimens from samples retrieved from two distinctive palaeoenvironmental settings in the western Mediterranean Tethys: three sections from the Lusitanian Basin in Portugal and one section from the Causses Basin in southern France. For each specimen, coccolith length, width and central area length and width were measured using specific measurement software. Statistical and data analysis tools were used in order to evaluate significant resemblances and/or differences between different size groups and samples. Two main clusters are acknowledged from this study. Small-sized Lotharingius is <4μm and large-sized Lotharingius is >4μm. In addition to common evolutionary processes, changes in palaeoenvironmental conditions were taken into account to explain such a size distribution. Although background evolution pushed the Lotharingius genus toward larger sizes, palaeoenvironmental conditions seemed to play an important role in the Toarcian-early Aalenian morphological evolution of this taxon. The production of larger coccoliths profited from stable environmental conditions, whereas smaller morphotypes dominated during periods marked by stress. The transition from small to large Lotharingius was synchronous in the two investigated western Tethys basins at the Bifrons-Gradata (or Variabilis) ammonite zone boundary (middle Toarcian), thus representing an important event for the Lower Jurassic calcareous nannofossil biostratigraphy.

Evidence for Strain-Specific Exometabolomic Responses of the Coccolithophore Emiliania huxleyi to Grazing by the Dinoflagellate Oxyrrhis marina

The coccolithophore Emiliania huxleyi forms massive blooms and plays a critical role in global elemental cycles, sequestering significant amounts of atmospheric carbon dioxide on geological time scales via production of calcium carbonate coccoliths and emitting dimethyl sulfoniopropionate (DMSP) which has the potential for increasing atmospheric albedo. Because grazing in pelagic systems is a major top-down force structuring microbial communities, the influence of grazers on E. huxleyi populations has been of interest to researchers. Roles of DMSP (and related metabolites) in interactions between E. huxleyi and protist grazers have been investigated, however, little is known about the release of other metabolites that may influence, or be influenced by, such grazing interactions. We used high-resolution mass spectrometry in an untargeted approach to survey the suite of low molecular weight compounds released by four different E. huxleyi strains in response to grazing by the dinoflagellate Oxyrrhis marina. Overall, a strikingly small number of metabolites were detected from E. huxleyi and O. marina cells, but these were distinctly informative to construct metabolic footprints. At most, E. huxleyi strains shared 25% of released metabolites. Furthermore, there appeared to be no unified metabolic response in E. huxleyi strains to grazing; rather these responses were strain specific. Concentrations of several metabolites also positively correlated with grazer activities, including grazing, ingestion, and growth rates; however, no single metabolite responded uniformly across all strains of E. huxleyi tested. Regardless, grazing clearly transformed the constituents of dissolved organic matter produced by these marine microbes. This study addresses several technical challenges, and presents a platform to further study the influence of chemical cues in aquatic systems and demonstrates the impact of strain diversity and grazing on the complexity of dissolved organic matter in marine systems.

Losses, Expansions, and Novel Subunit Discovery of Adaptor Protein Complexes in Haptophyte Algae

The phylum Haptophyta (Diaphoratickes) contains marine algae that perform biomineralization, extruding large, distinctive calcium carbonate scales (coccoliths) that completely cover the cell. Coccolith production is an important part of global carbon cycling; however, the membrane trafficking pathway by which they are secreted has not yet been elucidated. In most eukaryotes, post-Golgi membrane trafficking involves five heterotetrameric adaptor protein (AP) complexes, which impart cargo selection specificity. To better understand coccolith secretion, we performed comparative genomic, phylogenetic, and transcriptomic analyses of the AP complexes in Emiliania huxleyi strains 92A, Van556, EH2, and CCMP1516, and related haptophytes Gephyrocapsa oceanica and Isochrysis galbana; the latter has lost the ability to biomineralize. We show that haptophytes have a modified membrane trafficking system (MTS), as we found both AP subunit losses and duplications. Additionally, we identified a single conserved subunit of the AP-related TSET complex, whose expression suggests a functional role in membrane trafficking. Finally, we detected novel alpha adaptin ear and gamma adaptin ear proteins, the first of their kind to be described outside of opisthokonts. These novel ear proteins and the sculpting of the MTS may support the capacity for biomineralization in haptophytes, enhancing their ability to perform this highly specialized form of secretion.

Bacterial Diversity Associated with the Coccolithophorid Algae Emiliania huxleyi and Coccolithus pelagicus f. braarudii.

Coccolithophores are unicellular calcifying marine phytoplankton that can form large and conspicuous blooms in the oceans and make significant contributions to oceanic carbon cycling and atmospheric CO2 regulation. Despite their importance, the bacterial diversity associated with these algae has not been explored for ecological or biotechnological reasons. Bacterial membership of Emiliania huxleyi and Coccolithus pelagicus f. braarudii cultures was assessed using cultivation and cultivation-independent methods. The communities were species rich compared to other phytoplankton cultures. Community analysis identified specific taxa which cooccur in all cultures (Marinobacter and Marivita). Hydrocarbon-degrading bacteria were found in all cultures. The presence of Acidobacteria, Acidimicrobidae, Schlegelella, and Thermomonas was unprecedented but were potentially explained by calcification associated with coccolith production. One strain of Acidobacteria was cultivated and is closely related to a marine Acidobacteria isolated from a sponge. From this assessment of the bacterial diversity of coccolithophores, a number of biotechnological opportunities are evident, from bioprospecting for novel taxa such as Acidobacteria to helping understand the relationship between obligate hydrocarbonoclastic bacteria occurrence with phytoplankton and to revealing bacterial taxa that have a specific association with algae and may be suitable candidates as a means to improve the efficiency of mass algal cultivation.

Investigation of Cell Growth and Chlorophyll a Content of the Coccolithophorid Alga Emiliania huxleyi by Using Simple Bench-Top Flow Cytometry

The coccolithophorid alga Emiliania huxleyi produces micro-structured calcite particles, which are called coccoliths. Due to their unique and sophisticated structure, coccoliths are highly promising for different industrial applications, such as paper manufacturing, color and lacquer preparation. The mass production of coccoliths requires the evaluation of optimum cultivation conditions. This study investigates the impact of varying irradiance (10-1500 μmol m-2 s-1) on growth and chlorophyll a content of two calcifying strains CCMP371 and RCC1216 as well as on the non-calcifying strain RCC1217 (haploid form of RCC1217). The light kinetics contradicts the popular opinion, that E. huxleyi is an extraordinarily light tolerating alga in general. Photoinhibition was already observed at irradiance >500 μmol m-2 s-1 in the case of the calcifying strains. Furthermore, light requirements to grow at maximum growth rate, as well as thresholds towards photoinhibition were considerably different between calcifying and non-calcifying strains. The haplont required significantly higher irradiance to reach maximum μspec (>200 μmol m-2 s-1), while being much more tolerant to towards photoinhibition, which occurred not until 800 μmol m-2 s-1. Furthermore, a novel method was proposed to allow for the estimation of chlorophyll a content from flow cytometry data. By comprising an Advanced Fluorescence Ratio (AFLR), which considers culture heterogeneity, this method enables for simple chlorophyll a estimation also in older cultures of calcifying Emiliania huxleyi, which tend to build agglomerates.