DISCOVERY 2010: Spatial and temporal variability in a dynamic polar ecosystem

The Scotia Sea has been a focus of biological- and physical oceanographic study since the Discovery expeditions in the early 1900s. It is a physically energetic region with some of the highest levels of productivity in the Southern Ocean. It is also a region within which there have been greater than average levels of change in upper water column temperature. We describe the results of three cruises transecting the central Scotia Sea from south to north in consecutive years and covering spring, summer and autumn periods. We also report on some community level syntheses using both current-day and historical data from this region. A wide range of parameters were measured during the field campaigns, covering the physical oceanography of the region, air–sea CO2 fluxes, macro- and micronutrient concentrations, the composition and biomass of the nano-, micro- and mesoplankton communities, and the distribution and biomass of Antarctic krill and mesopelagic fish. Process studies examined the effect of iron-stress on the physiology of primary producers, reproduction and egestion in Antarctic krill and the transfer of stable isotopes between trophic layers, from primary consumers up to birds and seals. Community level syntheses included an examination of the biomass-spectra, food-web modelling, spatial analysis of multiple trophic layers and historical species distributions. The spatial analyses in particular identified two distinct community types: a northern warmer water community and a southern cold community, their boundary being broadly consistent with the position of the Southern Antarctic Circumpolar Current Front (SACCF). Temperature and ice cover appeared to be the dominant, over-riding factors in driving this pattern. Extensive phytoplankton blooms were a major feature of the surveys, and were persistent in areas such as South Georgia. In situ and bioassay measurements emphasised the important role of iron inputs as facilitators of these blooms. Based on seasonal DIC deficits, the South Georgia bloom was found to contain the strongest seasonal carbon uptake in the ice-free zone of the Southern Ocean. The surveys also encountered low-production, iron-limited regions, a situation more typical of the wider Southern Ocean. The response of primary and secondary consumers to spatial and temporal heterogeneity in production was complex. Many of the life-cycles of small pelagic organisms showed a close coupling to the seasonal cycle of food availability. For instance, Antarctic krill showed a dependence on early, non-ice-associated blooms to facilitate early reproduction. Strategies to buffer against environmental variability were also examined, such as the prevalence of multiyear life-cycles and variability in energy storage levels. Such traits were seen to influence the way in which Scotia Sea communities were structured, with biomass levels in the larger size classes being higher than in other ocean regions. Seasonal development also altered trophic function, with the trophic level of higher predators increasing through the course of the year as additional predator-prey interactions emerged in the lower trophic levels. Finally, our studies re-emphasised the role that the simple phytoplankton-krill-higher predator food chain plays in this Southern Ocean region, particularly south of the SACCF. To the north, alternative food chains, such as those involving copepods, macrozooplankton and mesopelagic fish, were increasingly important. Continued ocean warming in this region is likely to increase the prevalence of such alternative such food chains with Antarctic krill predicted to move southwards.

Particle export during a bloom of Emiliania huxleyi in the North-West European continental margin

Coccolithophores, the dominant pelagic calcifiers in the oceans, play a key role in the marine carbon cycle through calcification, primary production and carbon export, the main drivers of the biological CO2 pump. In May 2002 a cruise was conducted on the outer shelf of the North-West European continental margin, from the north Bay of Biscay to the Celtic Sea (47.0 degrees-50.5 degrees N, 5.0 degrees-11.0 degrees W), an area where massive blooms of Emiliania huxleyi are observed annually. Biogeochemical variables including primary production, calcification, partial pressure of CO2 (pCO(2)), chlorophyll-a (Chl-a), particle load, particulate organic and inorganic carbon (POC, PIC) and Th-234, were measured in surface waters to assess particle dynamic and carbon export in relation to the development of a coccolithophore bloom. We observed a marked northward decrease in Chl-a concentration and calcification rates: the bloom exhibited lower values and may be less well developed in the Goban Spur area. The export fluxes of POC and PIC from the top 80 m, determined using the ratios of POC and PIC to Th-234 of particles, ranged from 81 to 323 mg C m(-2) d(-1) and from 30 to 84 mg C m(-2) d(-1), respectively. The highest fluxes were observed in waters presenting a well-developed coccolithophore bloom, as shown by high reflectance of surface waters. This experiment confirms that the occurrence of coccolithophores promotes efficient export of organic and inorganic carbon on the North-West European margin.

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.

Volcanic ash fuels anomalous plankton bloom in subarctic northeast Pacific

Using multiple lines of evidence, we demonstrate that volcanic ash deposition in August 2008 initiated one of the largest phytoplankton blooms observed in the subarctic North Pacific. Unusually widespread transport from a volcanic eruption in the Aleutian Islands, Alaska deposited ash over much of the subarctic NE Pacific, followed by large increases in satellite chlorophyll. Surface ocean pCO2, pH, and fluorescence reveal that the bloom started a few days after ashfall. Ship-based measurements showed increased dominance by diatoms. This evidence points toward fertilization of this normally iron-limited region by ash, a relatively new mechanism proposed for iron supply to the ocean. The observations do not support other possible mechanisms. Extrapolation of the pCO2 data to the area of the bloom suggests a modest ∼0.01 Pg carbon export from this event, implying that even large-scale iron fertilization at an optimum time of year is not very efficient at sequestering atmospheric CO2.

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.

Coccolithophore bloom detection in the north east Atlantic using SeaWiFS: Algorithm description, application and sensitivity analysis

Coccolithophores are the largest source of calcium carbonate in the oceans and are considered to play an important role in oceanic carbon cycles. Current methods to detect the presence of coccolithophore blooms from Earth observation data often produce high numbers of false positives in shelf seas and coastal zones due to the spectral similarity between coccolithophores and other suspended particulates. Current methods are therefore unable to characterise the bloom events in shelf seas and coastal zones, despite the importance of these phytoplankton in the global carbon cycle. A novel approach to detect the presence of coccolithophore blooms from Earth observation data is presented. The method builds upon previous optical work and uses a statistical framework to combine spectral, spatial and temporal information to produce maps of coccolithophore bloom extent. Validation and verification results for an area of the north east Atlantic are presented using an in situ database (N = 432) and all available SeaWiFS data for 2003 and 2004. Verification results show that the approach produces a temporal seasonal signal consistent with biological studies of these phytoplankton. Validation using the in situ coccolithophore cell count database shows a high correct recognition rate of 80% and a low false-positive rate of 0.14 (in comparison to 63% and 0.34 respectively for the established, purely spectral approach). To guide its broader use, a full sensitivity analysis for the algorithm parameters is presented.

Reduction in photosystem II efficiency during a virus-controlled Emiliania huxleyi bloom

During viral infection of Emiliania huxleyi, laboratory studies have shown that photo-system (PS) II efficiency declines during the days post-infection and is thought to be associated with viral-induced interruption of electron transport rates between photosystems. However,measuring the impact of viral infection on PSII function in E. huxleyi populations from natural,taxonomically diverse phytoplankton communities is difficult, and whether this phenomenon occurs in nature is presently unknown. Here, chlorophyll fluorescence analysis was used to assess changes in PSII efficiency throughout an E. huxleyi bloom during a mesocosm experiment off the coast of Norway. Specifically, we aimed to determine whether a measurable suppression of the efficiency of PSII photochemistry could be observed due to viral infection of the natural E. huxleyi populations. During the major infection period prior to bloom collapse, there was a significant reduction in PSII efficiency with an average decrease in maximum PSII photochemical efficiency (Fv/Fm) of 17% and a corresponding 75% increase in maximum PSII effective absorption cross section(σPSII); this was concurrent with a significant decrease in E. huxleyi growth rates and an increase in E. huxleyi virus (EhV) production. As E. huxleyi populations dominated the phytoplankton community and potentially contributed up to 100% of the chlorophyll a pool, we believe that the variable chlorophyll fluorescence signal measured during this period was derived predominantly from E. huxleyi and, thus, reflects changes occurring within E. huxleyi cells. This is the first demonstration of suppression of PSII photochemistry occurring during viral infection of natural coccolithophore populations.