From silk to satellite: half a century of ocean colour anomalies in the Northeast Atlantic

Changes in phytoplankton dynamics influence marine biogeochemical cycles, climate processes, and food webs, with substantial social and economic consequences. Large-scale estimation of phytoplankton biomass was possible via ocean colour measurements from two remote sensing satellites – the Coastal Zone Color Scanner (CZCS, 1979-1986) and the Sea-viewing Wide Field-of-view Sensor (SeaWiFS, 1998-2010). Due to the large gap between the two satellite eras and differences in sensor characteristics, comparison of the absolute values retrieved from the two instruments remains challenging. Using a unique in situ ocean colour dataset that spans more than half a century, the two satellite-derived chlorophyll-a (Chl-a) eras are linked to assess concurrent changes in phytoplankton variability and bloom timing over the Northeast Atlantic Ocean and North Sea. Results from this unique re-analysis reflect a clear increasing pattern of Chl-a, a merging of the two seasonal phytoplankton blooms producing a longer growing season and higher seasonal biomass, since the mid-1980s. The broader climate plays a key role in Chl-a variability as the ocean colour anomalies parallel the oscillations of the Northern Hemisphere Temperature (NHT) since 1948.

Development and validation of a molecular technique for the analysis of archived formalin-preserved phytoplankton samples permits retrospective assessment of Emiliania huxleyi communities

The coccolithophores, particularly the species Emiliania huxleyi (Lohmann) Hay & Mohler, account for the bulk of global calcium carbonate production and as such play a fundamental role in global CO2 cycling and the carbonate chemistry of the oceans. To evaluate the response of this functional group to the effects of climate change, we undertook a feasibility study to determine whether a retrospective approach could be used on archived coccolithophore datasets. We demonstrate for the first time a technique for the extraction of E. huxleyi nucleic acids from archived formalin-fixed samples of the long-term Continuous Plankton Recorder. Molecular analysis of a nine year old formalin-fixed sample reveals the presence of a diverse population of E. huxleyi genotypes within a developing coccolithophore bloom. In addition, E. huxleyi sequences were amplified from a number of formalin-fixed samples, the earliest of which was collected in August 1972. This molecular assay promises the possibility of studying global variations in the distribution and genetic make-up of E. huxleyi communities over extensive periods of time. (c) 2008 Elsevier B.V. All rights reserved.

Growth and production of Pacific ocean perch (Sebastes alutus) in nursery habitats of the Gulf of Alaska

Nursery areas for juvenile fishes are often important for determining recruitment in marine populations by providing habitats that can maximize growth and thereby minimize mortality. Pacific ocean perch (POP, Sebastes alutus) have an extended juvenile period where they inhabit rocky nursery habitats. We examined POP nursery areas to link growth potential to recruitment. Juvenile POP were captured from nursery areas in 2004 and 2008, and estimated growth rates ranged from −0.19 to 0.60 g day−1 based on differences in size between June and August. Predicted growth rates from a bioenergetics model ranged from 0.05 to 0.49 g day−1 and were not significantly different than observed. Substrate preferences and the distribution of their preferred habitats were utilized to predict the extent of juvenile POP nursery habitat in the Gulf of Alaska. Based on densities of fish observed on underwater video transects and the spatial extent of nursery areas, we predicted 278 and 290 million juvenile POP were produced in 2004 and 2008. Growth potential for juvenile POP was reconstructed using the bioenergetics model, spring zooplankton bloom timing and duration and bottom water temperature for 1982–2008. When a single outlying recruitment year in 1986 was removed, growth potential experienced by juvenile POP in nursery areas was significantly correlated to the recruitment time-series from the stock assessment, explaining ∼30% of the variability. This research highlights the potential to predict recruitment using habitat-based methods and provides a potential mechanism for explaining some of the POP recruitment variability observed for this population.

Air/sea DMS gas transfer in the North Atlantic: evidence for limited interfacial gas exchange at high wind speed

Shipboard measurements of eddy covariance dimethylsulfide (DMS) air–sea fluxes and seawater concentration were carried out in the North Atlantic bloom region in June/July 2011. Gas transfer coefficients (k660) show a linear dependence on mean horizontal wind speed at wind speeds up to 11 m s−1. At higher wind speeds the relationship between k660 and wind speed weakens. At high winds, measured DMS fluxes were lower than predicted based on the linear relationship between wind speed and interfacial stress extrapolated from low to intermediate wind speeds. In contrast, the transfer coefficient for sensible heat did not exhibit this effect. The apparent suppression of air–sea gas flux at higher wind speeds appears to be related to sea state, as determined from shipboard wave measurements. These observations are consistent with the idea that long waves suppress near-surface water-side turbulence, and decrease interfacial gas transfer. This effect may be more easily observed for DMS than for less soluble gases, such as CO2, because the air–sea exchange of DMS is controlled by interfacial rather than bubble-mediated gas transfer under high wind speed conditions.

Dip in the gene pool: metagenomic survey of natural coccolithovirus communities

Despite the global oceanic distribution and recognized biogeochemical impact of coccolithoviruses (EhV), their diversity remains poorly understood. Here we employed a metagenomic approach to study the occurrence and progression of natural EhV community genomic variability. Analysis of EhV metagenomes from the early and late stages of an induced bloom led to three main discoveries. First, we observed resilient and specific genomic signatures in the EhV community associated with the Norwegian coast, which reinforce the existence of limitations to the capacity of dispersal and genomic exchange among EhV populations. Second, we identified a hyper-variable region (approximately 21 kbp long) in the coccolithovirus genome. Third, we observed a clear trend for EhV relative amino-acid diversity to reduce from early to late stages of the bloom. This study validated two new methodological combinations, and proved very useful in the discovery of new genomic features associated with coccolithovirus natural communities.

Satellite discrimination of Karenia mikimotoi and Phaeocystis harmful algal blooms in European coastal waters: Merged classification of ocean colour data

The detection of dense harmful algal blooms (HABs) by satellite remote sensing is usually based on analysis of chlorophyll-a as a proxy. However, this approach does not provide information about the potential harm of bloom, nor can it identify the dominant species. The developed HAB risk classification method employs a fully automatic data-driven approach to identify key characteristics of water leaving radiances and derived quantities, and to classify pixels into “harmful”, “non-harmful” and “no bloom” categories using Linear Discriminant Analysis (LDA). Discrimination accuracy is increased through the use of spectral ratios of water leaving radiances, absorption and backscattering. To reduce the false alarm rate the data that cannot be reliably classified are automatically labelled as “unknown”. This method can be trained on different HAB species or extended to new sensors and then applied to generate independent HAB risk maps; these can be fused with other sensors to fill gaps or improve spatial or temporal resolution. The HAB discrimination technique has obtained accurate results on MODIS and MERIS data, correctly identifying 89% of Phaeocystis globosa HABs in the southern North Sea and 88% of Karenia mikimotoi blooms in the Western English Channel. A linear transformation of the ocean colour discriminants is used to estimate harmful cell counts, demonstrating greater accuracy than if based on chlorophyll-a; this will facilitate its integration into a HAB early warning system operating in the southern North Sea.

Feeding and overwintering of Antarctic krill across its major habitats: The role of sea ice cover, water depth, and phytoplankton abundance

Antarctic krill (Euphausia superba) were sampled in contrasting habitats: a seasonally ice-covered deep ocean (Lazarev Sea), ice-free shelves at their northern range (South Georgia) and the Antarctic Peninsula (Bransfield Strait), and shelf and oceanic sites in the Scotia Sea. Across 92 stations, representing a year-round average, the food volume in krill stomachs comprised 71 +/- 29% algae, 17 +/- 21% protozoans, and 12 +/- 25% metazoans. Fatty acid trophic markers showed that copepods were consistently part of krill diet, not a switch food. In open waters, both diatom and copepod consumption increased with phytoplankton abundance. Under sea ice, ingestion of diatoms became rare, whereas feeding on copepods remained constant. During winter, larvae contained high but variable proportions of diatom markers, whereas in postlarvae the role of copepods increased with krill body length. Overwintering differed according to habitat. Krill from South Georgia had lower lipid stores than those from the Bransfield Strait or Lazarev Sea. Feeding effort was much reduced in Lazarev Sea krill, whereas most individuals from the Bransfield Strait and South Georgia contained phytoplankton and seabed detritus in their stomachs. Their retention of essential body reserves indicates that krill experienced most winter hardship in the Lazarev Sea, followed by South Georgia and then Bransfield Strait. This was reflected in the delayed development from juveniles to adults in the Lazarev Sea. Circumpolar comparisons of length frequencies suggest that krill growth conditions are more favorable in the southwest Atlantic than in the Lazarev Sea or off East Antarctica because of longer phytoplankton bloom periods and rewarding access to benthic food.

Ocean Net Heat Flux Influences Seasonal to Interannual Patterns of Plankton Abundance

Changes in the net heat flux (NHF) into the ocean have profound impacts on global climate. We analyse a long-term plankton time-series and show that the NHF is a critical indicator of ecosystem dynamics. We show that phytoplankton abundance and diversity patterns are tightly bounded by the switches between negative and positive NHF over an annual cycle. Zooplankton increase before the transition to positive NHF in the spring but are constrained by the negative NHF switch in autumn. By contrast bacterial diversity is decoupled from either NHF switch, but is inversely correlated (r=-0.920) with the magnitude of the NHF. We show that the NHF is a robust mechanistic tool for predicting climate change indicators such as spring phytoplankton bloom timing and length of the growing season.

Operational monitoring and forecasting of bathing water quality through exploiting satellite Earth observation and models: The AlgaRisk demonstration service

Coastal zones and shelf-seas are important for tourism, commercial fishing and aquaculture. As a result the importance of good water quality within these regions to support life is recognised worldwide and a number of international directives for monitoring them now exist. This paper describes the AlgaRisk water quality monitoring demonstration service that was developed and operated for the UK Environment Agency in response to the microbiological monitoring needs within the revised European Union Bathing Waters Directive. The AlgaRisk approach used satellite Earth observation to provide a near-real time monitoring of microbiological water quality and a series of nested operational models (atmospheric and hydrodynamic-ecosystem) provided a forecast capability. For the period of the demonstration service (2008–2013) all monitoring and forecast datasets were processed in near-real time on a daily basis and disseminated through a dedicated web portal, with extracted data automatically emailed to agency staff. Near-real time data processing was achieved using a series of supercomputers and an Open Grid approach. The novel web portal and java-based viewer enabled users to visualise and interrogate current and historical data. The system description, the algorithms employed and example results focussing on a case study of an incidence of the harmful algal bloom Karenia mikimotoi are presented. Recommendations and the potential exploitation of web services for future water quality monitoring services are discussed.

Dimethylsulfide gas transfer coefficients from algal blooms in the Southern Ocean

Air–sea dimethylsulfide (DMS) fluxes and bulk air–sea gradients were measured over the Southern Ocean in February–March 2012 during the Surface Ocean Aerosol Production (SOAP) study. The cruise encountered three distinct phytoplankton bloom regions, consisting of two blooms with moderate DMS levels, and a high biomass, dinoflagellate-dominated bloom with high seawater DMS levels (> 15 nM). Gas transfer coefficients were considerably scattered at wind speeds above 5 m/s. Bin averaging the data resulted in a linear relationship between wind speed and mean gas transfer velocity consistent with that previously observed. However, the wind-speed-binned gas transfer data distribution at all wind speeds is positively skewed. The flux and seawater DMS distributions were also positively skewed, which suggests that eddy covariance-derived gas transfer velocities are consistently influenced by additional, log-normal noise. A flux footprint analysis was conducted during a transect into the prevailing wind and through elevated DMS levels in the dinoflagellate bloom. Accounting for the temporal/spatial separation between flux and seawater concentration significantly reduces the scatter in computed transfer velocity. The SOAP gas transfer velocity data show no obvious modification of the gas transfer–wind speed relationship by biological activity or waves. This study highlights the challenges associated with eddy covariance gas transfer measurements in biologically active and heterogeneous bloom environments.

Trait-based representation of diatom functional diversity in a plankton functional type model of the eutrophied southern North Sea

We introduce a trait-based description of diatom functional diversity to an existing plankton functional type (PFT) model, implemented for the eutrophied coastal ecosystem in the Southern Bight of the North Sea. The trait-based description represents a continuum of diatom species, each characterized by a distinct cell volume, and includes size dependence of four diatom traits: the maximum growth rate, the half-saturation constants for nutrient uptake, the photosynthetic efficiency, and the relative affinity of copepods for diatoms. Through competition under seasonally varying forcing, the fitness of each diatom varies throughout time, and the outcome of competition results in a changing community structure. The predicted seasonal change in mean cell volume of the community is supported by field observations: smaller diatoms, which are more competitive in terms of resource acquisition, prevail during the first spring bloom, whereas the summer bloom is dominated by larger species which better resist grazing. The size-based model is used to determine the ecological niche of diatoms in the area and identifies a range of viable sizes that matches observations. The general trade-off between small, competitive diatoms and large, grazing-resistant species is a convenient framework to study patterns in diatom functional diversity. PFT models and trait-based approaches constitute promising complementary tools to study community structure in marine ecosystems.

Ocean acidification with (de)eutrophication will alter future phytoplankton growth and succession.

Human activity causes ocean acidification (OA) though the dissolution of anthropogenically generated CO2 into seawater, and eutrophication through the addition of inorganic nutrients. Eutrophication increases the phytoplankton biomass that can be supported during a bloom, and the resultant uptake of dissolved inorganic carbon during photosynthesis increases water-column pH (bloom-induced basification). This increased pH can adversely affect plankton growth. With OA, basification commences at a lower pH. Using experimental analyses of the growth of three contrasting phytoplankton under different pH scenarios, coupled with mathematical models describing growth and death as functions of pH and nutrient status, we show how different conditions of pH modify the scope for competitive interactions between phytoplankton species. We then use the models previously configured against experimental data to explore how the commencement of bloom-induced basification at lower pH with OA, and operating against a background of changing patterns in nutrient loads, may modify phytoplankton growth and competition. We conclude that OA and changed nutrient supply into shelf seas with eutrophication or de-eutrophication (the latter owing to pollution control) has clear scope to alter phytoplankton succession, thus affecting future trophic dynamics and impacting both biogeochemical cycling and fisheries.

Drivers and effects of Karenia mikimotoi blooms in the western English Channel

Naturally occurring red tides and harmful algal blooms (HABs) are of increasing importance in the coastal environment and can have dramatic effects on coastal benthic and epipelagic communities worldwide. Such blooms are often unpredictable, irregular or of short duration, and thus determining the underlying driving factors is problematic. The dinoflagellate Karenia mikimotoi is an HAB, commonly found in the western English Channel and thought to be responsible for occasional mass finfish and benthic mortalities. We analysed a 19-year coastal time series of phytoplankton biomass to examine the seasonality and interannual variability of K. mikimotoi in the western English Channel and determine both the primary environmental drivers of these blooms as well as the effects on phytoplankton productivity and oxygen conditions. We observed high variability in timing and magnitude of K. mikimotoi blooms, with abundances reaching >1000 cells mL�1 at 10 m depth, inducing up to a 12-fold increase in the phytoplankton carbon content of the water column. No long-term trends in the timing or magnitude of K. mikimotoi abundance were evident from the data. Key driving factors were identified as persistent summertime rainfall and the resultant input of low-salinity high-nutrient river water. The largest bloom in 2009 was associated with highest annual primary production and led to considerable oxygen depletion at depth, most likely as a result of enhanced biological breakdown of bloom material; however, this oxygen depletion may not affect zooplankton. Our data suggests that K. mikimotoi blooms are not only a key and consistent feature of western English Channel productivity, but importantly can potentially be predicted from knowledge of rainfall or river discharge.

A puzzle with many pieces: the genetic structure and diversity of Phaeocystis antarctica Karsten (Prymnesiophyta)

Strong ocean current systems characterize the Southern Ocean. The genetic structure of marine phytoplankton species is believed to depend mainly on currents. Genetic estimates of the relatedness of populations of phytoplankton species therefore should provide a proxy showing to what extent different geographic regions are interconnected by the ocean current systems. In this study, spatial and temporal patterns of genetic diversity were studied in the circumpolar prymnesiophyte Phaeocystis antarctica Karsten using seven nuclear microsatellite loci. Analyses were conducted for 86 P. antarctica isolates sampled around the Antarctic continent between 1982 and2007. The resultsrevealed highgenetic diversity without singlegenotypes recurringeven amongisolateswithin a bloom or originating from the same bucket of water. Populations of P. antarctica were significantly differentiated among the oceanic regions. However, some geographically distant populations were more closely related to each other than they were to other geographically close populations. Temporal haplotype turnover within regions was also suggested by the multilocus fingerprints. Our data suggest that even within blooms of P. antarctica genetic diversity and population sizes are large but exchange between different regions canbe limited. Positive and significant inbreeding coefficients hint at further regional substructure of populations, suggestingthat patches, once isolated from one another, may not reconnect. These data emphasize that even for planktonic species in a marine ecosystem that is influenced by strong currents, significant breaks in geneflow may occur.

On the ultrastructure of Gephyrocapsa oceanica (Haptophyta) life stages.

Gephyrocapsa oceanica is a cosmopolitan bloom-forming coccolithophore species belonging to the haptophyte order Isochrysidales and family Noëlaerhabdaceae. Exclusively pelagic, G. oceanica is commonly found in modern oceans and in fossil assemblages. Its sister species Emiliania huxleyi is known to possess a haplo-diplontic life cycle, the non-motile diploid coccolith-bearing cells alternating with haploid cells that are motile and covered by non-mineralized organic scales. Since the cytology and ultrastructure of other members of the Noëlaerhabdaceae has never been reported, it is not clear whether these features are common to the family. Here, we report on the ultrastructure of both the non-motile calcifying stage and the non-calcifying motile stage of G. oceanica. We found no significant ultrastructural differences between E. huxleyi and G. oceanica either in the calcifying diploid stage or the haploid phase. The similarities between these two morphospecies demonstrated a high degree of conservation of cytological features. We discuss the significance of these results in the light of the evolution of the Noelaerhabdaceae.