Coccolithophore Bloom Research Articles

Winter coccolithophore blooms in the Black Sea: Interannual variability and driving factors

The spatial and temporal variability of winter coccolithophore blooms in the Black Sea is studied using satellite measurements for 2003–2017 and Bio-Argo buoys data. Analysis shows that such blooms can occupy the entire upper 60-m layer with satellite-derived estimates of surface cell concentration reaching in some years 3.0·106 cells l−1. On average, winter coccolithophore blooms start in November–December, peak in the first decade of January, and terminate in February. They are characterized by significant interannual variability. Blooms in different years can be “basin-wide” covering most of the sea (2004–2005, 2005–2006, 2006–2007, 2012–2013) and “local” when they are observed only in certain areas of the Black Sea, most often in its southern part. A detailed analysis of the evolution of winter blooms has shown that they are usually triggered by intense storms, which are followed after 1–2 weeks by the initial growth of coccolithophores in the zone of storm-driven upwelling. Cross-shelf nutrients exchange may contribute to the coccolithophore growth over the continental slope. Intense winter coccolithophore blooms are observed in years with low autumn concentrations of chlorophyll-a, i.e., during periods of weakening of the typical diatom blooming autumn-winter. The strongest correlation on annual time scales is found between bloom intensity and decrease of the autumn-winter wind curl. We suggest that low wind curl causes a weakening of the basin-scale cyclonic circulation and intensifies the cross-shelf exchange, impacting the Black Sea trophic structure in the preceding summer period and may promote the winter bloom of coccolithophores in such years.

Vliyaniye sluchayev tsveteniya kokkolitoforid na bioopticheskiye kharakteristiki vod Chernogo morya po dannym nablyudeniy v 2012 i 2017 godakh

This paper presents and discusses the results of optical and biological measurements performed during the coccolithophore blooms. This study uses data of measurements from a stationary oceanographic platform (near the Southern coast of Crimea, 44°23' N, 33°59' E) in July 2012 (7–16 July) and in May 2017 (24–31 May), and data obtained during the R/V “Professor Vodyanitsky” expedition to the northern part of the Black Sea in June 2017 (14–30 June). Observation periods coincided with intensive blooms of coccolithophore. The measurements of the Secchi disk depth, light beam attenuation coefficient and reflectance coefficient, as well as biological determinations of the coccolithophore cells number in water samples, are considered and analyzed. In addition to field data, satellite information on the sea reflectance and backscattering are used. The beam attenuation coefficient, reflectance coefficient, backscattering coefficient and coccolithophore concentrations were increased as compared with no bloom period. The Secchi disk depth was abnormally low. The number of coccolithophore cells, according to biological data, varied from 150 thousand cells/L to 1.7 million cells/L and averaged 0.6 ± 0.4 million cells/L. According to these data, the estimated concentration of shed coccoliths was (1.46 ± 0.42) 1011 m-3 . The average coccolith concentrations calculated from satellite data were (1.29 ± 0.23) 1011 m-3 in July 2012, (1.32 ± 0.12) 1011 m-3 in May 2017, and (2.37 ± 0.69) 1011 m-3 in June 2017. During blooms, high variability was observed not only in optical properties, but also in biological properties, and the relationship between their spatial distribution was weak. From the optical point of view, the indicator of bloom is the number of coccoliths, not cells, which can lead to an incorrect description of the biological situation.

Bio-optical evidence for increasing Phaeocystis dominance in the Barents Sea

Increasing contributions of prymnesiophytes such as Phaeocystis pouchetii and Emiliania huxleyi to Barents Sea (BS) phytoplankton production have been suggested based on in situ observations of phytoplankton community composition, but the scattered and discontinuous nature of these records confounds simple inference of community change or its relationship to salient environmental variables. However, provided that meaningful assessments of phytoplankton community composition can be inferred based on their optical characteristics, ocean-colour records offer a potential means to develop a synthesis between sporadic in situ observations. Existing remote-sensing algorithms to retrieve phytoplankton functional types based on chlorophyll-a (chl-a) concentration or indices of pigment packaging may, however, fail to distinguish Phaeocystis from other blooms of phytoplankton with high pigment packaging, such as diatoms. We develop a novel algorithm to distinguish major phytoplankton functional types in the BS and apply it to the MODIS-Aqua ocean-colour record, to study changes in the composition of BS phytoplankton blooms in July, between 2002 and 2018, creating time series of the spatial distribution and intensity of coccolithophore, diatom and Phaeocystis blooms. We confirm a north-eastward expansion in coccolithophore bloom distribution, identified in previous studies, and suggest an inferred increase in chl-a concentrations, reported by previous researchers, may be partly explained by increasing frequencies of Phaeocystis blooms. This article is part of the theme issue 'The changing Arctic Ocean: consequences for biological communities, biogeochemical processes and ecosystem functioning'.

Effect of a Coccolithophore Bloom on the Underwater Light Field and the Albedo of the Water Column

The goal of this work is to study the influence of coccolithophore blooms on the underwater light field and albedo of the water column. A coccolithophore is a single-celled alga with spherical cells surrounded by disk-shaped calcite plates (coccolites), which produce strong light scattering. Because of that, we can observe coccolithophore blooms on satellite ocean color images. We calculated the angular underwater radiance distributions and their integral parameters by the exact numerical method with the input parameters, corresponding to real conditions observed in the Barents Sea and Black Sea. Using the results of the exact calculations, we estimated, for various situations, the accuracy of the approximating formulas applied to the assessment of the water radiance reflectance and the diffuse attenuation coefficients and we make recommendations for their application. As a finding of practical importance, we can note the estimate of the accuracy of the widely used Gordon’s formula for the diffuse attenuation coefficient; this formula results in large errors under strong coccolithophore blooms. We also mention the interesting and important results concerning the features of the asymptotic regime under such conditions.

Polarized lidar and ocean particles: insights from a mesoscale coccolithophore bloom.

Oceanographic lidar can provide remote estimates of the vertical distribution of suspended particles in natural waters, potentially revolutionizing our ability to characterize marine ecosystems and properly represent them in models of upper ocean biogeochemistry. However, lidar signals exhibit complex dependencies on water column inherent optical properties (IOPs) and instrument characteristics, which complicate efforts to derive meaningful biogeochemical properties from lidar return signals. In this study, we used a ship-based system to measure the lidar attenuation coefficient (α) and linear depolarization ratio (δ) across a variety of optically and biogeochemically distinct water masses, including turbid coastal waters, clear oligotrophic waters, and calcite rich waters associated with a mesoscale coccolithophore bloom. Sea surface IOPs were measured continuously while underway to characterize the response of α and δ to changes in particle abundance and composition. The magnitude of α was consistent with the diffuse attenuation coefficient (Kd), though the α versus Kd relationship was nonlinear. δ was positively related to the scattering optical depth and the calcite fraction of backscattering. A statistical fit to these data suggests that the polarized scattering properties of calcified particles are distinct and contribute to measurable differences in the lidar depolarization ratio. A better understanding of the polarized scattering properties of coccolithophores and other marine particles will further our ability to interpret polarized oceanographic lidar measurements and may lead to new techniques for measuring the material properties of marine particles remotely.

Simulation of factors affecting <i>Emiliania huxleyi</i> blooms in Arctic and sub-Arctic seas by CMIP5 climate models: model validation and selection

Abstract. The observed warming in the Arctic is more than double the global average, and this enhanced Arctic warming is projected to continue throughout the 21st century. This rapid warming has a wide range of impacts on polar and sub-polar marine ecosystems. One of the examples of such an impact on ecosystems is that of coccolithophores, particularly Emiliania huxleyi, which have expanded their range poleward during recent decades. The coccolithophore E. huxleyi plays an essential role in the global carbon cycle. Therefore, the assessment of future changes in coccolithophore blooms is very important. Currently, there are a large number of climate models that give projections for various oceanographic, meteorological, and biochemical variables in the Arctic. However, individual climate models can have large biases when compared to historical observations. The main goal of this research was to select an ensemble of climate models that most accurately reproduces the state of environmental variables that influence the coccolithophore E. huxleyi bloom over the historical period when compared to reanalysis data. We developed a novel approach for model selection to include a diverse set of measures of model skill including the spatial pattern of some variables, which had not previously been included in a model selection procedure. We applied this method to each of the Arctic and sub-Arctic seas in which E. huxleyi blooms have been observed. Once we have selected an optimal combination of climate models that most skilfully reproduce the factors which affect E. huxleyi, the projections of the future conditions in the Arctic from these models can be used to predict how E. huxleyi blooms will change in the future. Here, we present the validation of 34 CMIP5 (fifth phase of the Coupled Model Intercomparison Project) atmosphere–ocean general circulation models (GCMs) over the historical period 1979–2005. Furthermore, we propose a procedure of ranking and selecting these models based on the model's skill in reproducing 10 important oceanographic, meteorological, and biochemical variables in the Arctic and sub-Arctic seas. These factors include the concentration of nutrients (NO3, PO4, and SI), dissolved CO2 partial pressure (pCO2), pH, sea surface temperature (SST), salinity averaged over the top 30 m (SS30 m), 10 m wind speed (WS), ocean surface current speed (OCS), and surface downwelling shortwave radiation (SDSR). The validation of the GCMs' outputs against reanalysis data includes analysis of the interannual variability, seasonal cycle, spatial biases, and temporal trends of the simulated variables. In total, 60 combinations of models were selected for 10 variables over six study regions using the selection procedure we present here. The results show that there is neither a combination of models nor one model that has high skill in reproducing the regional climatic-relevant features of all combinations of the considered variables in target seas. Thereby, an individual subset of models was selected according to our model selection procedure for each combination of variable and Arctic or sub-Arctic sea. Following our selection procedure, the number of selected models in the individual subsets varied from 3 to 11. The paper presents a comparison of the selected model subsets and the full-model ensemble of all available CMIP5 models to reanalysis data. The selected subsets of models generally show a better performance than the full-model ensemble. Therefore, we conclude that within the task addressed in this study it is preferable to employ the model subsets determined through application of our procedure than the full-model ensemble.

Coccolithophore biodiversity controls carbonate export in the Southern Ocean

Abstract. Southern Ocean waters are projected to undergo profound changes in their physical and chemical properties in the coming decades. Coccolithophore blooms in the Southern Ocean are thought to account for a major fraction of the global marine calcium carbonate (CaCO3) production and export to the deep sea. Therefore, changes in the composition and abundance of Southern Ocean coccolithophore populations are likely to alter the marine carbon cycle, with feedbacks to the rate of global climate change. However, the contribution of coccolithophores to CaCO3 export in the Southern Ocean is uncertain, particularly in the circumpolar subantarctic zone that represents about half of the areal extent of the Southern Ocean and where coccolithophores are most abundant. Here, we present measurements of annual CaCO3 flux and quantitatively partition them amongst coccolithophore species and heterotrophic calcifiers at two sites representative of a large portion of the subantarctic zone. We find that coccolithophores account for a major fraction of the annual CaCO3 export, with the highest contributions in waters with low algal biomass accumulations. Notably, our analysis reveals that although Emiliania huxleyi is an important vector for CaCO3 export to the deep sea, less abundant but larger species account for most of the annual coccolithophore CaCO3 flux. This observation contrasts with the generally accepted notion that high particulate inorganic carbon accumulations during the austral summer in the subantarctic Southern Ocean are mainly caused by E. huxleyi blooms. It appears likely that the climate-induced migration of oceanic fronts will initially result in the poleward expansion of large coccolithophore species increasing CaCO3 production. However, subantarctic coccolithophore populations will eventually diminish as acidification overwhelms those changes. Overall, our analysis emphasizes the need for species-centred studies to improve our ability to project future changes in phytoplankton communities and their influence on marine biogeochemical cycles.

Phytoplankton Bloom and Photosynthetically Active Radiation in Coastal Waters

The effect of a phytoplankton bloom on the change with depth of photosynthetically available radiation (PAR), the spectral downwelling irradiance in coastal waters, and the phytoplankton ability to absorb PAR in the sea is estimated. The 10-fold increase in the chlorophyll a concentration (from 0.4 to 4.0 mg/m3) during a diatom/ dinoflagellate bloom leads to a decrease in water transparency and a narrowing of the photosynthesis zone. As a consequence, the average PAR within the phytoplankton habitat layer is almost halved. A diatom/dinoflagellate bloom is accompanied by changes in the downwelling irradiance spectrum at the same PAR level (optical depths). The penetrating irradiance maximum near the bottom of the euphotic zone shifts to longer wavelengths (~550–600 nm) as compared to background conditions (~500–550 nm). The increase in diatom/dinoflagellate biomass and the concomitant change in the downwelling irradiance spectrum are accompanied by a decrease in the specific absorption coefficient of phytoplankton pigments up to about three times that under background conditions at the same optical depths. The downwelling irradiance spectrum at fixed optical depths and the ability of cells to absorb light change slightly as compared to background conditions when coccolithophores bloom.

Seasonal dynamics of carbonate chemistry, nutrients and CO2 uptake in a sub-Arctic fjord

Environmental change can have a significant impact on biogeochemical cycles at high latitudes and be particularly important in ecologically valuable fjord ecosystems. Seasonality in biogeochemical cycling in a sub-Arctic fjord of northern Norway (Kaldfjorden) was investigated from October 2016 to September 2018. Monthly changes in total inorganic carbon (CT), alkalinity (AT), major nutrients and calcium carbonate saturation (Ω) were driven by freshwater discharge, biological production and mixing with subsurface carbon-rich coastal water. Stable oxygen isotope ratios indicated that meteoric water (snow melt, river runoff, precipitation) had stratified and freshened surface waters, contributing to 81% of the monthly CT deficit in the surface layer. The timing and magnitude of freshwater inputs played an important role in Ω variability, reducing AT and CT by dilution. This dilution effect was strongly counteracted by the opposing effect of primary production that dominated surface water Ω seasonality. The spring phytoplankton bloom rapidly depleted nitrate and CT to drive highest Ω (~2.3) in surface waters. Calcification reduced AT and CT, which accounted for 21% of the monthly decrease in Ω during a coccolithophore bloom. Freshwater runoff contributed CT, AT and silicates of terrestrial origin to the fjord. Lowest surface water Ω (~1.6) resulted from organic matter remineralisation and mixing into subsurface water during winter and spring. Surface waters were undersaturated with respect to atmospheric CO2, resulting in modest uptake of –0.32 ± 0.03 mol C m–2 yr–1. Net community production estimated from carbon drawdown was 14 ± 2 g C m–2 yr–1 during the productive season. Kaldfjorden currently functions as an atmospheric CO2 sink of 3.9 ± 0.3 g C m–2 yr–1. Time-series data are vital to better understand the processes and natural variability affecting biogeochemical cycling in dynamic coastal regions and thus better predict the impact of future changes on important fjord ecosystems.

Summer and winter coccolithophore blooms in the Black Sea and their impact on production of dissolved organic matter from Bio-Argo data

Bio-Argo measurements of the backscattering coefficient ( bbp ) were used to investigate the time-depth evolution of coccolithophore blooms in the Black Sea. Five years of Bio-Argo data obtained in 2014–2018 revealed two distinct peaks of bbp corresponding to the winter and early summer coccolithophore blooms. The latter started in the upper mixed layer (UML) in April–May and was characterized by the highest coccolithophore concentrations. During the most extensive summer bloom in 2017, its estimates reach 10 × 10 6 cells l −1 . The summer blooms occupied the UML (0–10 m) and a seasonal thermocline (10–30 m). The lower boundary of the bloom was related to the position of isopycnal 1014 kg m −3 , which deepens in May–July due to summer heating. Consequently, the coccolithophore bloom deepened to 20–30 m and terminated rapidly in July. Bloom termination was accompanied by a significant rise in light attenuation ( kd ) in the sea basin. This peak was attributed to the release of dissolved organic carbon (DOC), which was possibly related to viral lysis and the exudation of lipids from coccolithophore cells. Data on the kd was used to estimate the seasonal variability in DOC in the Black Sea. Maximal estimated values of DOC were observed at 15–35 m depth in June-August and coincided with the early summer coccolithophore bloom termination. The winter coccolithophore bloom started in October–November in the UML. The maximum bbp was observed in January. High values of bbp were observed down to a depth of 60 m during the maximal deepening of the mixed layer. The winter blooms were distinctly observed in MODIS satellite images, where they were characterized by high reflectance and relatively low chlorophyll concentrations. The estimated coccolithophore concentration in winter was lower than that in summer, but column-averaged bbp values were comparable. The winter coccolithophore bloom reached a peak within one month after the autumn peak of chlorophyll A, indicating the possible importance of the nutrients recycled after the diatom autumn bloom. In contrast to summer, the maximum DOC observed at the surface preceded the winter coccolithophore bloom, and the mass DOC production was probably attributed to the excretion and lysis of the non-calcified phytoplankton cells. • 5 year Bio-Argo data showed regular winter and summer coccolithophore blooms. • Summer bloom starts in May in the upper 20 m and deepen in June–July to 20–30 m. • Winter bloom starts in October–November and peaks in January, occupying 0-60 m layer. • Estimated column-integrated cell concentrations in winter and summer are comparable. • Peak of DOC is observed in August at 15–35 m depth during termination of summer bloom.

Estimation of seasonal and inter-annual variations of phytoplankton in the Black Sea on the base of remote sensed data procession and chlorophyll a in situ measurements

The patterns of the seasonal and inter annual variability in the development of phytoplankton, including coc- colithophores, were studied in the northeastern part of the Black Sea in various seasons. The research was carried out on the base of remote sensed data provided by the Modis Aqua ocean color scanner and long-term ship observations. The results indicate that, in the coastal and open waters of the Black Sea, the development of phytoplankton reaches the highest level in the fall-winter and winter-spring seasons, i.e., in the periods of active winter convection. It was noted that there is a relationship between the level of coccolithophore development in the summer period and the minimal temperature of the sea surface layer in the preceding winter. The most active coccolithophore blooms, were observed after the coldest winters. Unusual winter and early spring coccolithophore blooms taking place in mild hydro meteorological conditions were registered in the Black Sea, on the base of satellite data and were confirmed by in situ measurements.

Formation, Development, and Propagation of a Rare Coastal Coccolithophore Bloom

AbstractThis study examines an unprecedented bloom of Emiliania huxleyi along the California coast during the NE Pacific warm anomaly of 2014–2015. Observations of coccolithophore populations from microscopy and flow cytometry, surface current data derived from high‐frequency radar, and satellite ocean color imagery were used to track the population dynamics of the bloom in the Santa Barbara Channel. Results show a coastal bloom of mostly E. huxleyi that reached cell concentrations up to 5.7 × 106 cells per liter and a maximum spatial extent of 1,220 km2. We speculate that the rare cooccurrence of warm water, high water column stability, and an extensive preceding diatom bloom during the anomaly contributed to the development of this bloom. Flow cytometry measurements provided insight on the phases of bloom development (e.g., growth versus senescence) with calcified cells comprising up to 64% of particles containing chlorophyll a and detached‐coccolith:cell ratios ranging from 10 to >100. Lagrangian particle trajectories estimated during two nonoverlapping 48‐ and 72‐hr periods showed the changes in the surface structure of the bloom due to advection by surface currents and nonconservative biological and physical processes. Time rates of change of particulate inorganic carbon were estimated along particle trajectories, with rates ranging from −4 to 6 μmol·L−1·day−1. The approach presented here is likely to be useful for understanding the evolution of coastal phytoplankton bloom events in a general setting.

ИССЛЕДОВАНИЕ БИООПТИЧЕСКИХ ХАРАКТЕРИСТИК ВОД ПОВЕРХНОСТНОГО СЛОЯ БАРЕНЦЕВА И НОРВЕЖСКОГО МОРЕЙ ЛЕТОМ 2017 ГОДА

A study of the surface layer bio-optical properties of the Barents and Norwegian Seas in the summer of 2017 is carried out. Ship data were obtained during the 68th cruise of the R/V ‘Akademik Mstislav Keldysh’ (June-August 2017). Using a flow-through system, the fluorescence intensities of chlorophyll ‘a’ and dissolved organic matter as well as the salinity and temperature of the water surface layer along the ship’s route were continuously recorded. Seawater samples were taken for spectral fluorescence and absorbance measurements performed with a laser spectrometer and an integrating cavity absorption meter. The results are compared with the data of direct determinations of the chlorophyll concentration. Frequent continuous cloudiness prevented the use of ocean color data for the Norwegian Sea. In the coccolithophore bloom area in the Barents Sea, the results of shipboard measurements are compared with the data of OLCI satellite scanner. In this area, standard OLCI algorithms overestimate chlorophyll concentration, while the regression algorithm works better than based on neural networks. Comparison of the fluorescence and absorption spectra has shown the possibility of carrying out a rapid assessment of the chlorophyll concentration using optical methods. The change in the coefficients of the regression equations of chlorophyll fluorescence intensity and its concentration, determined by direct methods for different regions, is shown.

Seasonal and Interannual Variability of Phytoplankton in the Black Sea on the Basis of Remote Sensing Data and In Situ Measurements of Chlorophyll-a

The patterns of the seasonal and interannual variability in the development of phytoplankton, including coccolithophores, were studied in the northeastern part of the Black Sea in various seasons. The research was carried out on the basis of long-term observations and data provided by the Modis Aqua ocean color scanner. This paper reports that, in the coastal and open waters of the Black Sea, the development of phytoplankton reaches the highest level in the fall–winter and winter–spring seasons, i.e., in the periods of active winter convection. Coastal waters are also characterized by expressed spring peak of chlorophyll “a” concentrations associated with phytoplankton bloom in March. Winter and spring coccolithophore blooms, which are unusual for the Black Sea, were registered on the basis of satellite data. It was noted that there is a relationship between the level of coccolithophore development in the summer period and the minimal temperature of the sea surface layer in the preceding winter. The most active coccolithophore blooms, which were registered in 2006 and 2012, were observed after the coldest winters.

The influence of submesoscale eddies on hydrochemical parameters and structural and functional characteristics of phytoplankton in the north-eastern part of the Black Sea

On June 7, 2018, a sub-mesoscale anticyclonic eddy induced by the wind (north-east) was registered on the shelf in the area of the city of Gelendzhik. With the help of field multidisciplinary expedition ship surveys, it was shown that this eddy exists in the layer above the seasonal thermocline. At the periphery of the eddy weak variability of hydrochemical parameters and quantitative indicators of phytoplankton were recorded. The result of the formation of such eddy structure was a shift in the structure of phytoplankton – the annual observed coccolithophores bloom was not registered.

Spatial and Temporal Variability of Coccolithophore Blooms in the Eastern Bering Sea

AbstractCoccolithophores are a widespread group of marine phytoplankton that produce plates of calcium carbonate that cover their cells. Large blooms of coccolithophores may significantly influence the biogeochemical properties of the ocean and atmosphere and trophic dynamics of the marine ecosystem. Because of the important implications of coccolithophore blooms, their timely monitoring and reporting is necessary for ecosystem management. To communicate with ecosystem management stakeholders, we developed an annual Coccolithophore Bloom Index (CBI) for the eastern Bering Sea shelf using satellite ocean color data. Comparisons between in situ and satellite data and the CBI (years 1997–2017) were used to examine the hypotheses regarding environmental influences on interannual bloom variability. A significant nonlinear relationship with summer stratification was found: the CBI was higher during years with either very low or very high stratification. In addition, while the blooms usually occurred over the middle shelf (50‐ to 100‐m depth), more of the bloom was located over the shallow (30–50 m) inner shelf when stratification was low. Spatial correspondence between nutrient concentrations (nitrate and ammonium) and the areal extent of the coccolithophore bloom provides tantalizing but nonconclusive evidence that nutrient availability plays a role in bloom formation and location.

A 40-year global data set of visible-channel remote-sensing reflectances and coccolithophore bloom occurrence derived from the Advanced Very High Resolution Radiometer catalogue

Abstract. A consistently calibrated 40-year-long data set of visible-channel remote-sensing reflectance has been derived from the Advanced Very High Resolution Radiometer (AVHRR) sensor global time series. The data set uses as its source the Pathfinder Atmospheres – Extended (PATMOS-x) v5.3 Climate Data Record for top-of-atmosphere (TOA) visible-channel reflectances. This paper describes the theoretical basis for the atmospheric correction procedure and its subsequent implementation, including the necessary ancillary data files used and quality flags applied, in order to determine remote-sensing reflectance. The resulting data set is produced at daily, and archived at monthly, resolution, on a 0.1∘×0.1∘ grid at https://doi.org/10.1594/PANGAEA.892175. The primary aim of deriving this data set is to highlight regions of the global ocean affected by highly reflective blooms of the coccolithophorid Emiliania huxleyi (where lith concentration >2–5×104 mL−1) over the past 40 years.

Colin Brownlee.

Colin Brownlee.

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.

Expression profiling of host and virus during a coccolithophore bloom provides insights into the role of viral infection in promoting carbon export.

The cosmopolitan coccolithophore Emiliania huxleyi is a unicellular eukaryotic alga that forms vast blooms in the oceans impacting large biogeochemical cycles. These blooms are often terminated due to infection by the large dsDNA virus, E. huxleyi virus (EhV). It was recently established that EhV-induced modulation of E. huxleyi metabolism is a key factor for optimal viral infection cycle. Despite the huge ecological importance of this host-virus interaction, the ability to assess its spatial and temporal dynamics and its possible impact on nutrient fluxes is limited by current approaches that focus on quantification of viral abundance and biodiversity. Here, we applied a host and virus gene expression analysis as a sensitive tool to quantify the dynamics of this interaction during a natural E. huxleyi bloom in the North Atlantic. We used viral gene expression profiling as an index for the level of active infection and showed that the latter correlated with water column depth. Intriguingly, this suggests a possible sinking mechanism for removing infected cells as aggregates from the E. huxleyi population in the surface layer into deeper waters. Viral infection was also highly correlated with induction of host metabolic genes involved in host life cycle, sphingolipid, and antioxidant metabolism, providing evidence for modulation of host metabolism under natural conditions. The ability to track and quantify defined phases of infection by monitoring co-expression of viral and host genes, coupled with advance omics approaches, will enable a deeper understanding of the impact that viruses have on the environment.