Identification of senescence and death in Emiliania huxleyi and Thalassiosira pseudonana: Cell staining chlorophyll alterations and dimethylsulfoniopropionate (DMSP) metabolism

We measured membrane permeability, hydrolytic enzyme, and caspase-like activities using fluorescent cell stains to document changes caused by nutrient exhaustion in the coccolithophore Emiliania huxleyi and the diatom Thalassiosira pseudonana, during batch-culture nutrient limitation. We related these changes to cell death, pigment alteration, and concentrations of dimethylsulfide (DMS) and dimethylsulfoniopropionate (DMSP) to assess the transformation of these compounds as cell physiological condition changes. E. huxleyi persisted for 1 month in stationary phase; in contrast, T. pseudonana cells rapidly declined within 10 d of nutrient depletion. T. pseudonana progressively lost membrane integrity and the ability to metabolize 5-chloromethylfluorescein diacetate (CMFDA; hydrolytic activity), whereas E. huxleyi developed two distinct CMFDA populations and retained membrane integrity (SYTOX Green). Caspase-like activity appeared higher in E. huxleyi than in T. pseudonana during the post-growth phase, despite a lack of apparent mortality and cell lysis. Photosynthetic pigment degradation and transformation occurred in both species after growth; chlorophyll a (Chl a) degradation was characterized by an increase in the ratio of methoxy Chl a : Chl a in T. pseudonana but not in E. huxleyi, and the increase in this ratio preceded loss of membrane integrity. Total DMSP declined in T. pseudonana during cell death and DMS increased. In contrast, and in the absence of cell death, total DMSP and DMS increased in E. huxleyi. Our data show a novel chlorophyll alteration product associated with T. pseudonana death, suggesting a promising approach to discriminate nonviable cells in nature.

Dynamics of phytoplankton communities during late summer around the tip of the Antarctic Peninsula

The composition and distribution of phytoplankton assemblages around the tip of the Antarctic Peninsula were studied during two summer cruises (February/March 2008 and 2009). Water samples were collected for HPLC/CHEMTAX pigment and microscopic analysis. A great spatial variability in chlorophyll a (Chl a) was observed in the study area: highest levels in the vicinity of the James Ross Island (exceeding 7 mg m−3 in 2009), intermediate values (0.5 to 2 mg m−3) in the Bransfield Strait, and low concentrations in the Weddell Sea and Drake Passage (below 0.5 mg m−3). Phytoplankton assemblages were generally dominated by diatoms, especially at coastal stations with high Chl a concentration, where diatom contribution was above 90% of total Chl a. Nanoflagellates, such as cryptophytes and/or Phaeocystis antarctica, replaced diatoms in open-ocean areas (e.g., Weddell Sea). Many species of peridinin-lacking autotrophic dinoflagellates (e.g., Gymnodinium spp.) were also important to total Chl a biomass at well-stratified stations of Bransfield Strait. Generally, water column structure was the most important environmental factor determining phytoplankton communities’ biomass and distribution. The HPLC pigment data also allowed the assessment of different physiological responses of phytoplankton to ambient light variation. The present study provides new insights about the dynamics of phytoplankton in an undersampled region of the Southern Ocean highly susceptible to global climate change.

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.

Variation in ocean colour may help predict cod and haddock recruitment

Characteristics of the spring and fall phytoplankton blooms in spawning areas on the Scotian Shelf, Canada, were estimated from remote sensing data. These blooms, along with anomalies in the North Atlantic Oscillation, were used to explain variation in the recruitment of 4 populations of cod and haddock. We tested the effects of the timing of the bloom using the chlorophyll a (chl a) signal, the maximum amount of chl a, the timing of the diatom bloom, and the maximum relative dominance of diatoms on the recruitment (to Age 1) of cod and haddock on the Scotian Shelf. Models were run separately for the effects of the spring and fall blooms. Only 3 of 10 models tested (0-lag) explained significant (80 to 92%) variation in recruitment. However, the performance of these models was not consistent across populations or species, suggesting that generalities about how spring and fall phytoplankton blooms affect recruitment cannot yet be made. The differences among models suggest that fish larvae are probably adapted locally to food production and thus indirectly to the characteristics of the phytoplankton bloom, which in turn are influenced by regional (meso-scale) oceanographic conditions.

Variation among northeast Atlantic regions in the responses of zooplankton to climate change: Not all areas follow the same path

Broad scale climate forcing can interact with local environmental processes to affect the observed ecological phenomena. This causes potential problems of over-extrapolation for results from a limited number of sites or the averaging out of region-specific responses if data from too wide an area are combined. In this study, an area similar in extent to the Celtic Biscay Large Marine Ecosystem, but including off-shelf areas, was partitioned using clustering of satellite chlorophyll (chl-a) measurements. The resulting clusters were used to define areas over which to combine copepod data from the Continuous Plankton Recorder. Following filtering due to data limitations, nine regions were defined with sufficient records for analysis. These regions were consistent with known oceanographic structure in the study area. Off-shelf regions showed a progressively later timing in the seasonal peak of chl-a measurements moving northwards. Generalised additive models were used to estimate seasonal and multiannual signals in the adult and juvenile stages of Calanus finmarchicus, C. helgolandicus and the Paracalanus–Pseudocalanus group. Associations between variables (sea surface temperature (SST), phenology and annual abundance) differed among taxonomic groups, but even within taxonomic groups, relationships were not consistent across regions. For example, in the deep waters off Spain and Portugal the annual abundance of Calanus finmarchicus has a weak positive association with SST, in contrast to the pattern in most other regions. The regions defined in this study provide an objective basis for investigations into the long term dynamics of plankton populations and suggest suitable sub regions for deriving pelagic system indicators.

Non-linearities, regime shifts and recovery: The recent influence of climate on Black Sea chlorophyll

The Black Sea ecosystem experienced severe eutrophication-related degradation during the 1970s and 1980s. However, in recent years the Black Sea has shown some signs of recovery which are often attributed to a reduction in nutrient loading. Here, SeaWiFS chlorophyll a (chl a), a proxy for phytoplankton biomass, is used to investigate spatio-temporal patterns in Black Sea phytoplankton dynamics and to explore the potential role of climate in the Black Sea's recovery. Maps of chl a anomalies, calculated relative to the 8 year mean, emphasize spatial and temporal variability of phytoplankton biomass in the Black Sea, particularly between the riverine-influenced Northwest Shelf and the open Black Sea. Evolution of phytoplankton biomass has shown significant spatial variability of persistence of optimal bloom conditions between three major regions of the Black Sea. With the exception of 2001, chl a has generally decreased during our 8 year time-series. However, the winter of 2000–2001 was anomalously warm with low wind stress, resulting in reduced vertical mixing of the water column and retention of nutrients in the photic zone. These conditions were associated with anomalously high levels of chl a throughout much of the open Black Sea during the following spring and summer. The unusual climatic conditions occurring in 2001 may have triggered a shift in the Black Sea's chl a regime. The long-term significance of this recent shift is still uncertain but illustrates a non-linear response to climate forcing that makes future ecosystem changes in the pelagic Black Sea ecosystem difficult to predict.

A long-term chlorophyll dataset reveals regime shift in North Sea phytoplankton biomass unconnected to nutrient levels

During the 1980s, a rapid increase in the Phytoplankton Colour Index (PCI), a semiquantitative visual estimate of algal biomass, was observed in the North Sea as part of a regionwide regime shift. Two new data sets created from the relationship between the PCI and SeaWiFS chlorophyll a (Chl a) quantify differences in the previous and current regimes for both the anthropogenically affected coastal North Sea and the comparatively unaffected open North Sea. The new regime maintains a 13% higher Chl a concentration in the open North Sea and a 21% higher concentration in coastal North Sea waters. However, the current regime has lower total nitrogen and total phosphorus concentrations than the previous regime, although the molar N: P ratio in coastal waters is now well above the Redfield ratio and continually increasing. Besides becoming warmer, North Sea waters are also becoming clearer (i.e., less turbid), thereby allowing the normally light-limited coastal phytoplankton to more effectively utilize lower concentrations of nutrients. Linear regression analyses indicate that winter Secchi depth and sea surface temperature are the most important predictors of coastal Chl a, while Atlantic inflow is the best predictor of open Chl a; nutrient concentrations are not a significant predictor in either model. Thus, despite decreasing nutrient concentrations, Chl a continues to increase, suggesting that climatic variability and water transparency may be more important than nutrient concentrations to phytoplankton production at the scale of this study.

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.

Absorption-based algorithm of primary production for total and size-fractionated phytoplankton in coastal waters

Most satellite models of production have been designed and calibrated for use in the open ocean. Coastal waters are optically more complex, and the use of chlorophyll a (chl a) as a first-order predictor of primary production may lead to substantial errors due to significant quantities of coloured dissolved organic matter (CDOM) and total suspended material (TSM) within the first optical depth. We demonstrate the use of phytoplankton absorption as a proxy to estimate primary production in the coastal waters of the North Sea and Western English Channel for both total, micro- and nano+pico-phytoplankton production. The method is implemented to extrapolate the absorption coefficient of phytoplankton and production at the sea surface to depth to give integrated fields of total and micro- and nano+pico-phytoplankton primary production using the peak in absorption coefficient at red wavelengths. The model is accurate to 8% in the Western English Channel and 22% in this region and the North Sea. By comparison, the accuracy of similar chl a based production models was >250%. The applicability of the method to autonomous optical sensors and remotely sensed aircraft data in both coastal and estuarine environments is discussed.

Quantifying the impact of environmental variables upon catch-per-unit-effort of the Blue Shark Prionace glauca in the Western English Channel

The effect of environmental variables on blue shark Prionace glauca catch per unit effort (CPUE) in a recreational fishery in the western English Channel, between June and September 1998–2011, was quantified using generalized additive models (GAMs). Sea surface temperature (SST) explained 1·4% of GAM deviance, and highest CPUE occurred at 16·7° C, reflecting the optimal thermal preferences of this species. Surface chlorophyll a concentration (CHL) significantly affected CPUE and caused 27·5% of GAM deviance. Additionally, increasing CHL led to rising CPUE, probably due to higher productivity supporting greater prey biomass. The density of shelf-sea tidal mixing fronts explained 5% of GAM deviance, but was non-significant, with increasing front density negatively affecting CPUE. Time-lagged frontal density significantly affected CPUE, however, causing 12·6% of the deviance in a second GAM and displayed a positive correlation. This outcome suggested a delay between the evolution of frontal features and the subsequent accumulation of productivity and attraction of higher trophic level predators, such as P. glauca.

Quantifying the impact of environmental variables upon catch-per-unit-effort of the Blue Shark Prionace glauca in the Western English Channel

The effect of environmental variables on blue shark Prionace glauca catch per unit effort (CPUE) in a recreational fishery in the western English Channel, between June and September 1998–2011, was quantified using generalized additive models (GAMs). Sea surface temperature (SST) explained 1·4% of GAM deviance, and highest CPUE occurred at 16·7° C, reflecting the optimal thermal preferences of this species. Surface chlorophyll a concentration (CHL) significantly affected CPUE and caused 27·5% of GAM deviance. Additionally, increasing CHL led to rising CPUE, probably due to higher productivity supporting greater prey biomass. The density of shelf-sea tidal mixing fronts explained 5% of GAM deviance, but was non-significant, with increasing front density negatively affecting CPUE. Time-lagged frontal density significantly affected CPUE, however, causing 12·6% of the deviance in a second GAM and displayed a positive correlation. This outcome suggested a delay between the evolution of frontal features and the subsequent accumulation of productivity and attraction of higher trophic level predators, such as P. glauca.

On the temporal consistency of chlorophyll products derived from three ocean-colour sensors

Satellite ocean-colour sensors have life spans lasting typically five-to-ten years. Detection of long-term trends in chlorophyll-a concentration (Chl-a) using satellite ocean colour thus requires the combination of different ocean-colour missions with sufficient overlap to allow for cross-calibration. A further requirement is that the different sensors perform at a sufficient standard to capture seasonal and inter-annual fluctuations in ocean colour. For over eight years, the SeaWiFS, MODIS-Aqua and MERIS ocean-colour sensors operated in parallel. In this paper, we evaluate the temporal consistency in the monthly Chl-a time-series and in monthly inter-annual variations in Chl-a among these three sensors over the 2002–2010 time period. By subsampling the monthly Chl-a data from the three sensors consistently, we found that the Chl-a time-series and Chl-a anomalies among sensors were significantly correlated for >90% of the global ocean. These correlations were also relatively insensitive to the choice of three Chl-a algorithms and two atmospheric-correction algorithms. Furthermore, on the subsampled time-series, correlations between Chl-a and time, and correlations between Chl-a and physical variables (sea-surface temperature and sea-surface height) were not significantly different for >92% of the global ocean. The correlations in Chl-a and physical variables observed for all three sensors also reflect previous theories on coupling between physical processes and phytoplankton biomass. The results support the combining of Chl-a data from SeaWiFS, MODIS-Aqua and MERIS sensors, for use in long-term Chl-a trend analysis, and highlight the importance of accounting for differences in spatial sampling among sensors when combining ocean-colour observations.

Do oceanic loggerhead turtles Caretta caretta associate with oceanographic fronts? Evidence from the Canary Current Large Marine Ecosystem

ABSTRACT: Oceanographic fronts are physical interfaces between water masses that differ in properties such as temperature, salinity, turbidity and chl a enrichment. Bio-physical coupling along fronts can lead to the development of pelagic biodiversity hotspots. A diverse range of marine vertebrates have been shown to associate with fronts, using them as foraging and migration habitats. Elucidation of the ecological significance of fronts generates a better understanding of marine ecosystem functioning, conferring opportunities to improve management of anthropogenic activities in the oceans. This study presents novel insight into the oceanographic drivers of habitat use in a population of marine turtles characterised by an oceanic-neritic foraging dichotomy. Using satellite tracking data from adult female loggerhead turtles nesting at Cape Verde (n = 12), we test the hypothesis that oceanic-foraging loggerheads associate with mesocale (10s – to 100s of km) thermal fronts. We use high-resolution (1 km) composite front mapping to characterise frontal activity in the Canary Current Large Marine Ecosystem (LME) over 2 temporal scales: (1) seasonal front frequency and (2) 7-day front metrics. Our use-availability analysis indicates that oceanic loggerheads show a preference for the highly productive upwelling region between Cape Verde and mainland Africa, an area of intense frontal activity. Within the upwelling region, turtles appear to forage epipelagically around mesoscale thermal fronts, exploiting profitable foraging opportunities resulting from physical aggregation of prey.

Effect of phytoplankton size classes on bio-optical properties of phytoplankton in the Western Iberian coast: Application of models

Chlorophyll-a satellite products are routinely used in oceanography, providing a synoptic and global view of phytoplankton abundance. However, these products lack information on the community structure of the phytoplankton, which is crucial for ecological modelling and ecosystem studies. To assess the usefulness of existing methods to differentiate phytoplankton functional types (PFT) or phytoplankton size classes from satellite data, in-situ phytoplankton samples collected in the Western Iberian coast, on the North-East Atlantic, were analysed for pigments and absorption spectra. Water samples were collected in five different locations, four of which were located near the shore and another in an open-ocean, seamount region. Three different modelling approaches for deriving phytoplankton size classes were applied to the in situ data. Approaches tested provide phytoplankton size class information based on the input of pigments data (Brewin et al., 2010), absorption spectra data (Ciotti et al., 2002) or both (Uitz et al., 2008). Following Uitz et al. (2008), results revealed high variability in microphytoplankton chlorophyll-specific absorption coefficients, ranging from 0.01 to 0.09 m2 (mg chl)− 1 between 400 and 500 nm. This spectral analysis suggested, in one of the regions, the existence of small cells (< 20 μm) in the fraction of phytoplankton presumed to be microphytoplankton (based on diagnostic pigments). Ciotti et al. (2002) approach yielded the highest differences between modelled and measured absorption spectra for the locations where samples had high variability in community structure and cell size. The Brewin et al. (2010) pigment-based model was adjusted and a set of model coefficients are presented and recommended for future studies in offshore water of the Western Iberian coast.

Basking sharks and oceanographic fronts: quantifying associations in the north‐east Atlantic

Understanding the mechanisms linking oceanographic processes and marine vertebrate habitat use is critical to effective management of populations of conservation concern. The basking shark Cetorhinus maximus has been shown to associate with oceanographic fronts – physical interfaces at the transitions between water masses – to exploit foraging opportunities resulting from aggregation of zooplankton. However, the scale, significance and variability of these observed associations have not yet been established. Here, we quantify the influence of mesoscale (10s – 100s km) frontal activity on habitat use over timescales of weeks to months. We use animal-mounted archival tracking with composite front mapping via Earth Observation (EO) remote sensing to provide an oceanographic context to individual shark movements. We investigate levels of association with fronts occurring over two spatio-temporal scales, (i) broad-scale seasonally persistent frontal zones and (ii) contemporaneous mesoscale thermal and chl-a fronts. Using random walk simulations and logistic regression within an iterative generalised linear mixed modelling (GLMM) framework, we find that seasonal front frequency is a significant predictor of shark presence. Temporally-matched oceanographic metrics also indicate that sharks demonstrate a preference for productive regions, and associate with contemporaneous thermal and chl-a fronts more frequently than could be expected at random. Moreover, we highlight the importance of cross-frontal temperature change and persistence, which appear to interact to affect the degree of prey aggregation along thermal fronts. These insights have clear implications for understanding the preferred habitats of basking sharks in the context of anthropogenic threat management and marine spatial planning in the northeast Atlantic.