Distribution of calcifying and silicifying phytoplankton in relation to environmental and biogeochemical parameters during the late stages of the 2005 North East Atlantic Spring Bloom

The late stage of the North East Atlantic (NEA) spring bloom was investigated during June 2005 along a transect section from 45 to 66 degrees N between 15 and 20 degrees W in order to characterize the contribution of siliceous and calcareous phytoplankton groups and describe their distribution in relation to environmental factors. We measured several biogeochemical parameters such as nutrients, surface trace metals, algal pigments, biogenic silica (BSi), particulate inorganic carbon (PIC) or calcium carbonate, particulate organic carbon, nitrogen and phosphorus (POC, PON and POP, respectively), as well as transparent exopolymer particles (TEP). Results were compared with other studies undertaken in this area since the JGOFS NABE program. Characteristics of the spring bloom generally agreed well with the accepted scenario for the development of the autotrophic community. The NEA seasonal diatom bloom was in the late stages when we sampled the area and diatoms were constrained to the northern part of our transect, over the Icelandic Basin (IB) and Icelandic Shelf (IS). Coccolithophores dominated the phytoplankton community, with a large distribution over the Rockall-Hatton Plateau (RHP) and IB. The Porcupine Abyssal Plain (PAP) region at the southern end of our transect was the region with the lowest biomass, as demonstrated by very low Chla concentrations and a community dominated by picophytoplankton. Early depletion of dissolved silicic acid (DSi) and increased stratification of the surface layer most likely triggered the end of the diatom bloom, leading to coccolithophore dominance. The chronic Si deficiency observed in the NEA could be linked to moderate Fe limitation, which increases the efficiency of the Si pump. TEP closely mirrored the distribution of both biogenic silica at depth and prymnesiophytes in the surface layer suggesting the sedimentation of the diatom bloom in the form of aggregates, but the relative contribution of diatoms and coccolithophores to carbon export in this area still needs to be resolved.

Studies on growth rate and grazing mortality rate by microzooplankton of size-fractionated phytoplankton in spring and summer in the Jiaozhou Bay, China

Dilution experiments were performed to examine the growth rate and grazing mortality rate of size-fractionated phytoplankton at three typical stations, inside and outside the bay, in the spring and summer of 2003 in the Jiaozhou Bay, China. in spring, the phytoplankton community structure was similar among the three stations, and was mainly composed of nanophytoplankton, such as, Skeletonema costatum and Cylindrotheca closterium. The structure became significantly different for the three stations in summer, when the dominant species at Stas A, B and C were Chaetoceros curvisetus, Pseudo-nitzschia delicatissima, C. affinis, C. debilis, Coscinodiscus oculus-iridis and Paralia sulcata respectively. Tintinnopsis beroidea and T. tsingtaoensis were the dominant species in spring, whereas the microzooplankton was apparently dominated by Strombidium sp. in summer. Pico- and nanophytoplankton had a relatively greater growth rate than microzooplankton both in spring and summer. The growth rate and grazing mortality rate were 0.18 similar to 0.44 and 0.12 similar to 1.47 d(-1) for the total phytoplankton and 0.20 similar to 0.55 and 0.21 similar to 0.37 d-1 for nanophytoplankton in spring respectively. In summer, the growth rate and grazing mortality rate were 0.38 similar to 0.71 and 0.27 similar to 0.60 d-1 for the total phytoplankton and 0.11 similar to 1.18 and 0.41 similar to 0.72 d(-1) for nano- and microphytoplankton respectively. The carbon flux consumed by microzooplankton per day was 7.68 similar to 39.81 mg/m(3) in spring and 12.03 similar to 138.22 mg/m(3) in summer respectively. Microzooplankton ingested 17.56%similar to 92.19% of the phytoplankton standing stocks and 31.77%similar to 467.88% of the potential primary productivity in spring; in contrast, they ingested 34.60%similar to 83.04% of the phytoplankton standing stocks and 71.28%similar to 98.80% of the potential primary productivity in summer. Pico- and nanophytoplankton appeared to have relatively greater rates of growth and grazing mortality than microphytoplankton during the experimental period. The grazing rate of microzooplankton in summer was a little bit greater than that in spring because of the relatively higher incubation temperature and different dominant microzooplankton species. Microzooplankton preferred ingesting nanophytoplankton to microphytoplankton in spring, while they preferred ingesting picophytoplankton to nanophytoplankton and microphytoplankton in summer. Compared with the results of dilution experiments performed in various waters worldwide, the results are in the middle range.

Evolution of nutrient structure and phytoplankton composition in the Jiaozhou Bay ecosystem

The inventories of nutrients in the surface water and large phytoplankton( > 69 pm) were analyzed from the data set of JERS ecological database about a typical coastal waters, the Jiaozhou Bay, China, from 1960s for N, P and from 1980s; for Si. By examining long-term changes of nutrient concentration, calculating stoichiometric balance, and comparing diatom composition, Si limitation of diatom production was found to be more possible. The possibility of Si limitation was from 37% in 1980s to 50% in 1990s. Jiaozhou Bay ecosystem is becoming serious eutrophication, with notable increase of NO2-N, NO3-N and NH4-N from 0.1417 mumol/L, 0.5414 mumol/L, 1.7222 mumol/L in 1960s to 0.9551 mumol/L, 3.001 mumol/L, 8.0359 mumol/L in late 1990s respectively and prominent decrease of Si from 4.2614 mumol/L in 1980s to 1.5861 mumol/L in late 1990s; the nutrient structure is controlled by nitrogen; the main limiting nutrient is probably silicon; because of the Si limitation the phytoplankton community structure has changed drastically.

Preliminary study on different nutrient pools supplies for the phytoplankton growth in the Jiaozhou Bay in China in the fall of 2004

The source and significance. of two mitrients, nitrogen. and phosphorous, were investigated by a modified dilution method performed on seawater samples from the Jiaozhou Bay in autumn 2004. This modified dilution method accounted for the phytoplankton growth rate, microzooplankton grazing mortality rate, the external nutrient pools, as well as nutrient supplied through remineralization by microzooplankton. The results indicated that the phytoplankton net growth rate increased in turn from inside the bay, to outside the bay, to in the Xiaogang Harbor. The phytoplankton, maximum growth rates and microzooplankton grazing mortality rates were 1.14 and 0.92 d(-1) outside the bay, 0.42 and 0.32 d(-1) inside the bay and 0.98 and 0.62 d(-1) in the harbor respectively. Outside the bay, the remineralized nitrogen (K-r = 24.49) had heavy influence on the growth of the phytoplankton. Inside the bay, the remineralized phosphorus(K-r = 3.49) strongly affected the phytoplankton growth. In the harbor, the remineralized phosphorus (K-r = 3.73) was in larger demand by phytoplankton growth. The results demonstrated that the different nutrients pools supplied for phytoplankton growth were greatly in accordance with the phytoplankton community structure, microzooplankton grazing mortality rates and environmental conditions. It is revealed that, nutrient remineralization is much more important for the phytoplankton growth in the Jiaozhou Bay than previously believed.

Changes in nitrogen and phosphorus and their effects on phytoplankton in the Bohai Sea

Systematic studies of the changes in dissolved inorganic nitrogen (DIN) and dissolved inorganic phosphorus (DIP) and their effects on phytoplankton over the last 30 years in the Bohai Sea are presented. The amount of sewage disposal, use of fertilizer and the Huanghe River runoff were found to have a significant influence on the DIN or DIP concentrations in the Bohai Sea over the last 30 years. Moreover, the changes in DIN and DIP resulted in changes in the limiting nutrients of phytoplankton in the Bohai Sea from nitrogen in the early 1980s to nitrogen-phosphorus in the late 1980s, and then to phosphorus after the 1990s. In addition, changes in nitrogen and phosphorus had a significant effect on the phytoplankton community structure. The half saturation constant (K (s)) was used to evaluate the effect of nutrients on the phytoplankton community structure in the Bohai Sea over the last 30 years. Cell abundance percentages of dominant phytoplankton species with high K (s) values for phosphorus and low K (s) values for nitrogen have decreased since the 1980s, while those of dominant phytoplankton species with low K (s) values for phosphorus and high K (s) values for nitrogen increased during this period.

Decadal basin-scale changes in diatoms, dinoflagellates, and phytoplankton color across the North Atlantic

The Continuous Plankton Recorder (CPR) survey has been used to characterize phytoplankton and zooplankton space-time dynamics in the North Sea since 1931 and in the North Atlantic since 1939. Phytoplankton biomass is assessed from these samples by visual assessment of the green color of the silk mesh, the Phytoplankton Color Index (PCI), and the total count of diatoms and dinoflagellates. Species with a frequency of occurrence greater than 1% in the samples are used as indicator species of the community. We investigated (1) long-term fluctuations of phytoplankton biomass, total diatoms, and total dinoflagellates; (2) geographical variation of patterns; (3) the relationship between phytoplankton and climate forcing in the North Atlantic CPR samples; (4) the relative contribution of diatoms and dinoflagellates to the PCI; and (5) the fluctuations of the dominant species over the period of survey to provide more information on the processes linking climate to changes in the phytoplankton community. As a result of the differences in microscopic analysis methods prior to 1958, our analyses were conducted for the period ranging from 1958 to 2002. The North Atlantic was divided into six regions identified through bathymetric criteria and separated along a North-South axis. Based on 12 monthly time series, we demonstrate increasing trends in PCI and total dinoflagellates and a decrease in total diatoms.

Space-borne study of seasonal, multi-year, and decadal phytoplankton dynamics in the Bay of Biscay

Seasonal and inter-annual variations in phytoplankton community abundance in the Bay of Biscay are studied. Preliminarily processed by the National Aeronautics and Space Administration (NASA) to yield normalized water-leaving radiance and the top-of-the-atmosphere solar radiance, Sea-viewing Wide Field-of-View Sensor (SeaWiFS), Moderate Resolution Imaging Spectroradiometer (MODIS), and Coastal Zone Color Scanner (CZCS) data are further supplied to our dedicated retrieval algorithms to infer the sought for parameters. By applying the National Oceanic and Atmospheric Administration's (NOAA's) Advanced Very High Resolution Radiometer (AVHRR) data, the surface reflection coefficient in the only band in the visible spectrum is derived and employed for analysis. Decadal bridged time series of variations of diatom-dominated phytoplankton and green dinoflagellate Lepidodinium chlorophorum within the shelf zone and the coccolithophore Emiliania huxleyi in the pelagic area of the Bay are documented and analysed in terms of impacts of some biogeochemical and geophysical forcing factors.

The global distribution of phytoplankton size spectrum and size classes from their light-absorption spectra derived from satellite data

The absorption spectra of phytoplankton in the visible domain hold implicit information on the phytoplankton community structure. Here we use this information to retrieve quantitative information on phytoplankton size structure by developing a novel method to compute the exponent of an assumed power-law for their particle-size spectrum. This quantity, in combination with total chlorophyll-a concentration, can be used to estimate the fractional concentration of chlorophyll in any arbitrarily-defined size class of phytoplankton. We further define and derive expressions for two distinct measures of cell size of mixed. populations, namely, the average spherical diameter of a bio-optically equivalent homogeneous population of cells of equal size, and the average equivalent spherical diameter of a population of cells that follow a power-law particle-size distribution. The method relies on measurements of two quantities of a phytoplankton sample: the concentration of chlorophyll-a, which is an operational index of phytoplankton biomass, and the total absorption coefficient of phytoplankton in the red peak of visible spectrum at 676 nm. A sensitivity analysis confirms that the relative errors in the estimates of the exponent of particle size spectra are reasonably low. The exponents of phytoplankton size spectra, estimated for a large set of in situ data from a variety of oceanic environments (similar to 2400 samples), are within a reasonable range; and the estimated fractions of chlorophyll in pico-, nano- and micro-phytoplankton are generally consistent with those obtained by an independent, indirect method based on diagnostic pigments determined using high-performance liquid chromatography. The estimates of cell size for in situ samples dominated by different phytoplankton types (diatoms, prymnesiophytes, Prochlorococcus, other cyanobacteria and green algae) yield nominal sizes consistent with the taxonomic classification. To estimate the same quantities from satellite-derived ocean-colour data, we combine our method with algorithms for obtaining inherent optical properties from remote sensing. The spatial distribution of the size-spectrum exponent and the chlorophyll fractions of pico-, nano- and micro-phytoplankton estimated from satellite remote sensing are in agreement with the current understanding of the biogeography of phytoplankton functional types in the global oceans. This study contributes to our understanding of the distribution and time evolution of phytoplankton size structure in the global oceans.

The response of phytoplankton to climate variability associated with the North Atlantic Oscillation

Inter-annual variability in the timing of phytoplankton spring bloom and phytoplankton community structure in the central North Atlantic Ocean was quantified using ocean color data and continuous plankton recorder (CPR) data. This variability was related to the North Atlantic Oscillation using correlation analysis and multivariate auto-regression models. The initiation of the spring bloom derived from CPR phytoplankton color index data is similar to that derived from satellite chlorophyll, and exhibits a nominal correlation with the sea surface temperature (SST) and the North Atlantic Oscillation (NAO). The extrapolated spring bloom timing suggested later initiation of blooms in the mid-1980s and earlier initiation of blooms in the 1990s. The climatological phytoplankton community structure in the central North Atlantic is dominated by diatoms, except for a shift in community composition favoring dinoflagellates in August. The ratio of diatoms to total phytoplankton abundance and the ratio of dinoflagellates to total phytoplankton abundance are both closely correlated with the NAO and SST. The extended time series of phytoplankton community structure between 1985 and 2009, deduced from the time series of SST and NAO over the same interval, showed a decadal shift away from diatoms towards dinoflagellates. The linkages between the NAO, and changes in stratification and phytoplankton processes occur over a larger scale than previously observed.

Role of light and photophysiological properties on phytoplankton succession during the spring bloom in the north-western Mediterranean Sea

This study aimed to determine the role of light on the succession of the phytoplankton community during the spring bloom in the northwestern Mediterranean Sea. To this end, three successive Lagrangian experiments were carried out between March and April 2003. The three experiments correspond to distinct phases of the bloom development (pre-bloom, bloom peak and post-bloom, respectively) and therefore to different trophic conditions. Phytoplankton (sampled on a daily scale) was grouped in size-based classes (pico and nano+micro) each of them were characterised in terms of chemotaxonomic composition, primary production and photophysiological properties. The phytoplankton community evolved with time changing in both size-class dominance and specie/group dominance within each size class. The bloom peak was characterised by highly dynamic condition (i.e. vertical mixing) and by the dominance of both small (pico) and large (nano and micro) diatoms, as a result of their capacity to photoacclimate to changing light regimes (‘physiological plasticity’). Concluding, we suggest that the physiological adaptation to light is the main factor driving the succession of the phytoplankton community during the first phases of the bloom (until the onset of thermal stratification) in the western Mediterranean Sea.

Latitudinal phytoplankton distribution and the neutral theory of biodiversity

Aim Recent studies have suggested that global diatom distributions are not limited by dispersal, in the case of both extant species and fossil species, but rather that environmental filtering explains their spatial patterns. Hubbell's neutral theory of biodiversity provides a framework in which to test these alternatives. Our aim is to test whether the structure of marine phytoplankton (diatoms, dinoflagellates and coccolithophores) assemblages across the Atlantic agrees with neutral theory predictions. We asked: (1) whether intersite variance in phytoplankton diversity is explained predominantly by dispersal limitation or by environmental conditions; and (2) whether species abundance distributions are consistent with those expected by the neutral model. Location Meridional transect of the Atlantic (50 degrees N50 degrees S). Methods We estimated the relative contributions of environmental factors and geographic distance to phytoplankton composition using similarity matrices, Mantel tests and variation partitioning of the species composition based upon canonical ordination methods. We compared the species abundance distribution of phytoplankton with the neutral model using Etienne's maximum-likelihood inference method. Results Phytoplankton communities are slightly more determined by niche segregation (24%), than by dispersal limitation and ecological drift (17%). In 60% of communities, the assumption of neutrality in species' abundance distributions could not be rejected. In tropical zones, where oceanic gyres enclose large stable water masses, most communities showed low species immigration rates; in contrast, we infer that communities in temperate areas, out of oligotrophic gyres, have higher rates of species immigration. Conclusions Phytoplankton community structure is consistent with partial niche assembly and partial dispersal and drift assembly (neutral processes). The role of dispersal limitation is almost as important as habitat filtering, a fact that has been largely overlooked in previous studies. Furthermore, the polewards increase in immigration rates of species that we have discovered is probably caused by water mixing conditions and productivity.

Modelling a light-driven phytoplankton succession

We used a numerical model to investigate if and to what extent cellular photoprotective capacity accounts for succession and vertical distribution of marine phytoplankton species/groups. A model describing xanthophyll photoprotective activity in phytoplankton has been implemented in the European Regional Sea Ecosystem Model and applied at the station L4 in the Western English Channel. Primary producers were subdivided into three phytoplankton functional types defined in terms of their capacity to acclimate to different light-specific environments: low light (LL-type), high light (HL-type) and variable light (VL-type) adapted species. The LL-type is assumed to have low cellular level of xanthophyll-cycling pigments (PX) relative to the modelled photosynthetically active pigments (chlorophyll and fucoxanthin (FUCO) = PSP). The HL-type has high PX content relative to PSP while VL-type presents an intermediate PX to PSP ratio. Furthermore, the VL-type is capable of reversibly converting FUCO to PX and synthesizing new PX under high-light stress. In order to reproduce phytoplankton community succession with each of the three groups being dominant in different periods of the year, we had also to assume reduced grazing pressure on HL-adapted species. Model simulations realistically reproduce the observed seasonal patterns of pigments and nutrients highlighting the reasonability of the underpinning assumptions. Our model suggests that pigment-mediated photophysiology plays a primary role in determining the evolution of marine phytoplankton communities in the winter-spring period corresponding to the shoaling of the mixed layer and the increase of light intensity. Grazing selectivity however contributes to the phytoplankton community composition in summer.

Lagrangian evolution of DMS during the Southern Ocean gas exchange experiment: The effects of vertical mixing and biological community shift

Concentrations of dimethylsulfide (DMS) and its precursor dimethylsulfoniopropionate (DMSP) are highly variable in time and space. What is driving the variability in DMS(P), and can those variability be explained by physical processes and changes in the biological community? During the Southern Ocean Gas Exchange Experiment (SO GasEx) in the austral fall of 2008, two 3He/SF6 labeled patches were created in the surface water. SF6 and DMS were surveyed continuously in a Lagrangian framework, while direct measurements of air-sea exchange further constrained the gas budgets. Turbulent diffusivity at the base of the mixed layer was estimated from SF6 profiles and used to calculate the vertical fluxes of DMS and nutrients. Increasing mixed layer nutrient concentrations due to mixing were associated with a shift in the phytoplankton community structure, which in turned likely affected the sulfur dynamics on timescales of days. DMS concentration as well as air-sea DMS flux appeared to be decoupled from the DMSP concentration, possibly due to grazing and bacterial DMS production. Contrary to expectations, in an environment with high winds and modest productivity, physical processes (air-sea exchange, photochemistry, vertical mixing) only accounted for a small fraction of DMS loss from the surface water. Among the DMS sinks, inferred biological consumption most likely dominated during SO GasEx.

Strong responses of Southern Ocean phytoplankton communities to volcanic ash

Volcanic eruptions have been hypothesized as an iron supply mechanism for phytoplankton blooms; however, little direct evidence of stimulatory responses has been obtained in the field. Here we present the results of twenty-one 1–2 day bottle enrichment experiments from cruises in the South Atlantic and Southern Ocean which conclusively demonstrated a photophysiological and biomass stimulation of phytoplankton communities following supply of basaltic or rhyolitic volcanic ash. Furthermore, experiments in the Southern Ocean demonstrated significant phytoplankton community responses to volcanic ash supply in the absence of responses to addition of dissolved iron alone. At these sites, dissolved manganese concentrations were among the lowest ever measured in seawater, and we therefore suggest that the enhanced response to ash may have been a result of the relief of manganese (co)limitation. Our results imply that volcanic ash deposition events could trigger extensive phytoplankton blooms, potentially capable of significant impacts on regional carbon cycling.

Mechanisms shaping size structure and functional diversity of phytoplankton communities in the ocean

The factors regulating phytoplankton community composition play a crucial role in structuring aquatic food webs. However, consensus is still lacking about the mechanisms underlying the observed biogeographical differences in cell size composition of phytoplankton communities. Here we use a trait-based model to disentangle these mechanisms in two contrasting regions of the Atlantic Ocean. In our model, the phytoplankton community can self-assemble based on a trade-off emerging from relationships between cell size and (1) nutrient uptake, (2) zooplankton grazing, and (3) phytoplankton sinking. Grazing 'pushes' the community towards larger cell sizes, whereas nutrient uptake and sinking 'pull' the community towards smaller cell sizes. We find that the stable environmental conditions of the tropics strongly balance these forces leading to persistently small cell sizes and reduced size diversity. In contrast, the seasonality of the temperate region causes the community to regularly reorganize via shifts in species composition and to exhibit, on average, bigger cell sizes and higher size diversity than in the tropics. Our results raise the importance of environmental variability as a key structuring mechanism of plankton communities in the ocean and call for a reassessment of the current understanding of phytoplankton diversity patterns across latitudinal gradients.