Identification of a diagnostic marker to detect freshwater cyanophages of filamentous cyanobacteria

Cyanophages are viruses that infect the cyanobacteria, globally important photosynthetic microorganisms. Cyanophages are considered significant components of microbial communities, playing major roles in influencing host community diversity and primary productivity, terminating cyanobacterial water blooms, and influencing biogeochemical cycles. Cyanophages are ubiquitous in both marine and freshwater systems; however, the majority of molecular research has been biased toward the study of marine cyanophages. In this study, a diagnostic probe was developed to detect freshwater cyanophages in natural waters. Oligonucleotide PCR-based primers were designed to specifically amplify the major capsid protein gene from previously characterized freshwater cyanomyoviruses that are infectious to the filamentous, nitrogen-fixing cyanobacterial genera Anabaena and Nostoc. The primers were also successful in yielding PCR products from mixed virus communities concentrated from water samples collected from freshwater lakes in the United Kingdom. The probes are thought to provide a useful tool for the investigation of cyanophage diversity in freshwater environments.

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.

Changes in pH at the exterior surface of plankton with ocean acidification

Anthropogenically released CO2 is dissolving in the ocean, causing a decrease in bulk-seawater pH (ocean acidification). Projections indicate that the pH will drop 0.3 units from its present value by 2100 (ref. 1). However, it is unclear how the growth of plankton is likely to respond. Using simulations we demonstrate how pH and carbonate chemistry at the exterior surface of marine organisms deviates increasingly from those of the bulk sea water as organism metabolic activity and size increases. These deviations will increase in the future as the buffering capacity of sea water decreases with decreased pH and as metabolic activity increases with raised seawater temperatures. We show that many marine plankton will experience pH conditions completely outside their recent historical range. However, ocean acidification is likely to have differing impacts on plankton physiology as taxon-specific differences in organism size, metabolic activity and growth rates during blooms result in very different microenvironments around the organism. This is an important consideration for future studies in ocean acidification as the carbonate chemistry experienced by most planktonic organisms will probably be considerably different from that measured in bulk-seawater samples. An understanding of these deviations will assist interpretation of the impacts of ocean acidification on plankton of different size and metabolic activity.

Structure and diversity of intertidal benthic diatom assemblages in contrasting shores: a case study from the Tagus estuary1

The structure of intertidal benthic diatoms assemblages in the Tagus estuary was investigated during a 2-year survey, carried out in six stations with different sediment texture. Nonparametric multivariate analyses were used to characterize spatial and temporal patterns of the assemblages and to link them to the measured environmental variables. In addition, diversity and other features related to community physiognomy, such as size-class or life-form distributions, were used to describe the diatom assemblages. A total of 183 diatom taxa were identified during cell counts and their biovolume was determined. Differences between stations (analysis of similarity (ANOSIM), R=0.932) were more evident than temporal patterns (R=0.308) and mud content alone was the environmental variable most correlated to the biotic data (BEST, rho=0.863). Mudflat stations were typically colonized by low diversity diatom assemblages (H' similar to 1.9), mainly composed of medium-sized motile epipelic species (250-1,000 mu m(3)), that showed species-specific seasonal blooms (e.g., Navicula gregaria Donkin). Sandy stations had more complex and diverse diatom assemblages (H' similar to 3.2). They were mostly composed by a large set of minute epipsammic species (<250 mu m(3)) that, generally, did not show temporal patterns. The structure of intertidal diatom assemblages was largely defined by the interplay between epipelon and epipsammon, and its diversity was explained within the framework of the Intermediate Disturbance Hypothesis. However, the spatial distribution of epipelic and epipsammic life-forms showed that the definition of both functional groups should not be over-simplified.

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.

Coccolithophore surface distributions in the North Atlantic and their modulation of the air-sea flux of CO2 from 10 years of satellite Earth observation data

Coccolithophores are the primary oceanic phytoplankton responsible for the production of calcium carbonate (CaCO3). These climatically important plankton play a key role in the oceanic carbon cycle as a major contributor of carbon to the open ocean carbonate pump (similar to 50 %) and their calcification can affect the atmosphere-to-ocean (air-sea) uptake of carbon dioxide (CO2) through increasing the seawater partial pressure of CO2 (pCO(2)). Here we document variations in the areal extent of surface blooms of the globally important coccolithophore, Emiliania huxleyi, in the North Atlantic over a 10-year period (1998-2007), using Earth observation data from the Sea-viewing Wide Field-of-view Sensor (SeaWiFS). We calculate the annual mean sea surface areal coverage of E. huxleyi in the North Atlantic to be 474 000 +/- 104 000 km(2), which results in a net CaCO3 carbon (CaCO3-C) production of 0.14-1.71 Tg CaCO3-C per year. However, this surface coverage (and, thus, net production) can fluctuate inter-annually by -54/+81% about the mean value and is strongly correlated with the El Nino/Southern Oscillation (ENSO) climate oscillation index (r = 0.75, p < 0.02). Our analysis evaluates the spatial extent over which the E. huxleyi blooms in the North Atlantic can increase the pCO(2) and, thus, decrease the localised air-sea flux of atmospheric CO2. In regions where the blooms are prevalent, the average reduction in the monthly air-sea CO2 flux can reach 55%. The maximum reduction of the monthly air-sea CO2 flux in the time series is 155 %. This work suggests that the high variability, frequency and distribution of these calcifying plankton and their impact on pCO(2) should be considered if we are to fully understand the variability of the North Atlantic air-to-sea flux of CO2. We estimate that these blooms can reduce the annual N. Atlantic net sink atmospheric CO2 by between 3-28 %.

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.

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.

Patterns of jellyfish abundance in the North Atlantic

A number of explanations have been advanced to account for the increased frequency and intensity at which jellyfish (pelagic cnidarians and ctenophores) blooms are being observed, most of which have been locally directed. Here, we investigate seasonal and inter-annual patterns in abundance and distribution of jellyfish in the North Atlantic Ocean to determine if there have been any system-wide changes over the period 1946–2005, by analysing records of the presence of coelenterates from the Continuous Plankton Recorder (CPR) survey. Peaks in jellyfish abundance are strongly seasonal in both oceanic and shelf areas: oceanic populations have a mid-year peak that is more closely related to peaks in phyto- and zooplankton, whilst the later peak of shelf populations mirrors changes in SST and reflects processes of advection and aggregation. There have been large amplitude cycles in the abundance of oceanic and shelf jellyfish (although not synchronous) over the last 60 years, with a pronounced synchronous increase in abundance in both areas over the last 10 years. Inter-annual variations in jellyfish abundance in oceanic areas are related to zooplankton abundance and temperature changes, but not to the North Atlantic Oscillation or to a chlorophyll index. The long-term inter-annual abundance of jellyfish on the shelf could not be explained by any environmental variables investigated. As multi-decadal cycles and more recent increase in jellyfish were obvious in both oceanic and shelf areas, we conclude that these are likely to reflect an underlying climatic signal (and bottom-up control) rather than any change in fishing pressure (top-down control). Our results also highlight the role of the CPR data in investigating long-term changes in jellyfish, and suggest that the cnidarians sampled by the CPR are more likely to be holoplanktic hydrozoans and not the much larger meroplanktic scyphozoans as has been suggested previously.

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.

Changes in marine dinoflagellate and diatom abundance under climate change

Marine diatoms and dinoflagellates play a variety of key ecosystem roles as important primary producers (diatoms and some dinoflagellates) and grazers (some dinoflagellates). Additionally some are harmful algal bloom (HAB) species and there is widespread concern that HAB species may be increasing accompanied by major negative socio-economic impacts, including threats to human health and marine harvesting1, 2. Using 92,263 samples from the Continuous Plankton Recorder survey, we generated a 50-year (1960–2009) time series of diatom and dinoflagellate occurrence in the northeast Atlantic and North Sea. Dinoflagellates, including both HAB taxa (for example, Prorocentrum spp.) and non-HAB taxa (for example, Ceratium furca), have declined in abundance, particularly since 2006. In contrast, diatom abundance has not shown this decline with some common diatoms, including both HAB (for example, Pseudo-nitzschia spp.) and non-HAB (for example, Thalassiosira spp.) taxa, increasing in abundance. Overall these changes have led to a marked increase in the relative abundance of diatoms versus dinoflagellates. Our analyses, including Granger tests to identify criteria of causality, indicate that this switch is driven by an interaction effect of both increasing sea surface temperatures combined with increasingly windy conditions in summer.

Toxic marine microalgae and shellfish poisoning in the British isles: history, review of epidemiology, and future implications

The relationship between toxic marine microalgae species and climate change has become a high profile and well discussed topic in recent years, with research focusing on the possible future impacts of changing hydrological conditions on Harmful Algal Bloom (HAB) species around the world. However, there is very little literature concerning the epidemiology of these species on marine organisms and human health. Here, we examine the current state of toxic microalgae species around the UK, in two ways: first we describe the key toxic syndromes and gather together the disparate reported data on their epidemiology from UK records and monitoring procedures. Secondly, using NHS hospital admissions and GP records from Wales, we attempt to quantify the incidence of shellfish poisoning from an independent source. We show that within the UK, outbreaks of shellfish poisoning are rare but occurring on a yearly basis in different regions and affecting a diverse range of molluscan shellfish and other marine organisms. We also show that the abundance of a species does not necessarily correlate to the rate of toxic events. Based on routine hospital records, the numbers of shellfish poisonings in the UK are very low, but the identification of the toxin involved, or even a confirmation of a poisoning event is extremely difficult to diagnose. An effective shellfish monitoring system, which shuts down aquaculture sites when toxins exceed regularity limits, has clearly prevented serious impact to human health, and remains the only viable means of monitoring the potential threat to human health. However, the closure of these sites has an adverse economic impact, and the monitoring system does not include all toxic plankton. The possible geographic spreading of toxic microalgae species is therefore a concern, as warmer waters in the Atlantic could suit several species with southern biogeographical affinities enabling them to occupy the coastal regions of the UK, but which are not yet monitored or considered to be detrimental.

Effects of environmental conditions on the seasonal distribution of phytoplankton biomass in the North Sea

We study the spatial and seasonal variability of phytoplankton biomass (as phytoplankton color) in relation to the environmental conditions in the North Sea using data from the Continuous Plankton Recorder survey. By using only environmental fields and location as predictor variables we developed a nonparametric model (generalized additive model) to empirically explore how key environmental factors modulate the spatio-temporal patterns of the seasonal cycle of algal biomass as well as how these relate to the ,1988 North Sea regime shift. Solar radiation, as manifest through changes of sea surface temperature (SST), was a key factor not only in the seasonal cycle but also as a driver of the shift. The pronounced increase in SST and in wind speed after the 1980s resulted in an extension of the season favorable for phytoplankton growth. Nutrients appeared to be unimportant as explanatory variables for the observed spatio-temporal pattern, implying that they were not generally limiting factors. Under the new climatic regime the carrying capacity of the whole system has been increased and the southern North Sea, where the environmental changes have been more pronounced, reached a new maximum.

Sedimentation of acantharian cysts in the Iceland Basin: Strontium as a ballast for deep ocean particle flux, and implications for acantharian reproductive strategies

Acantharian cysts were discovered in sediment trap samples from spring 2007 at 2000 m in the Iceland Basin. Although these single-celled organisms contribute to particulate organic matter flux in the upper mesopelagic, their contribution to bathypelagic particle flux has previously been found negligible. Four time-series sediment traps were deployed and all collected acantharian cysts, which are reproductive structures. Across all traps, cysts contributed on average 3-22%, and 4―24% of particulate organic carbon and nitrogen (POC and PON) flux, respectively, during three separate collection intervals (the maximum contribution in any one trap was 48% for POC and 59% for PON). Strontium (Sr) flux during these 6 weeks reached 3 mg m―2 d―1. The acantharian celestite (SrSO4) skeleton clearly does not always dissolve in the mesopelagic as often thought, and their cysts can contribute significantly to particle flux at bathypelagic depths during specific flux events. Their large size (∼ I mm) and mineral ballast result in a sinking rate of ∼ 500 m d―1; hence, they reach the bathypelagic before dissolving. Our findings are consistent with a vertical profile of salinity-normalized Sr concentration in the Iceland Basin, which shows a maximum at 1700 m. Profiles of salinity-normalized Sr concentration in the subarctic Pacific reach maxima at ≤ 1500 m, suggesting that Acantharia might contribute to the bathypelagic particle flux there as well. We hypothesize that Acantharia at high latitudes use rapid, deep sedimentation of reproductive cysts during phytoplankton blooms so that juveniles can exploit the large quantity of organic matter that sinks rapidly to the deep sea following a bloom.