Long-term changes in abundance and distribution of microzooplankton in the NE Atlantic and North Sea

Long-term changes in mesozooplankton and phytoplankton populations have been well documented in the North Atlantic region, whereas data for microzooplankton are scarce. This neglected component of the plankton is a vital link in marine food-webs, grazing on smaller flagellates and cyanobacteria and in turn providing food for the larger mesozooplankton. We use the latest tintinnid (Ciliophora, Protista) data from the Continuous Plankton Recorder (CPR) survey in the NE Atlantic and North Sea to examine the phenology, distribution and abundance of this important group of ciliates. Presence/absence data came from 167 122 CPR samples collected between 1960 and 2009 and abundance data from 49 662 samples collected between 1996 and 2009. In the North Atlantic the genus Dictyocysta spp. dominated and Parafavella gigantea showed an increase in abundance around Iceland and Greenland. In the North Sea higher densities of Tintinnopsis spp., Favella serrata and Ptychocylis spp. were found. The presence of tintinnids in CPR samples collected in the North Atlantic has increased over the last 50 years and the seasonal window of high abundance has lengthened. Conversely in the North Sea there has been an overall reduction in abundance. We discuss possible drivers for these long-term changes and point the way forward to more holistic studies that examine how ecosystems, rather than just selected taxa, are responding to climate change.

Genetic analysis of Continuous Plankton Recorder (CPR) samples

Genetic analysis of Continuous Plankton Recorder (CPR) samples is enabling greater taxonomic resolution and the study of plankton population structure. Here, we present some results from the genetic analysis of CPR samples collected in the North Sea and north-eastern Atlantic that reveal the impacts of climate on benthic-pelagic coupling and the food web. We show that pronounced changes in the North Sea meroplankton are related to an increased abundance and spatial distribution of the larvae of the benthic echinoderm, Echinocardium cordatum. Key stages of reproduction in E. cordatum, gametogenesis and spawning, are influenced by winter and spring sea temperature (January-May). A stepwise increase in sea temperature after 1987, which has created warmer conditions earlier in the year, together with increased summer phytoplankton, may benefit the reproduction and survival of this benthic species. Competition between the larvae of E. cordatum and other holozooplanlcton taxa may now be altering the trophodynamics of the summer pelagic ecosystem. In the north-eastern Atlantic the genetic analysis of fish larvae sampled by the CPR has revealed an unprecedented increase in the abundance of juvenile snake pipefish, Entelurus aequoreiis since 2002. We argue that increased sea surface temperatures in winter and spring when the eggs of E. aqueoreus, which are brooded by the male, are developing and the young larvae are growing in the plankton are a likely cause. The increased abundance of this species in Atlantic and adjacent European seas already appears to be influencing the marine food web.

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.

The influence of ocean acidification on nitrogen regeneration and nitrous oxide production in the North-West European shelf sea

The assimilation and regeneration of dissolved inorganic nitrogen, and the concentration of N2O, was investigated at stations located in the NW European shelf sea during June/July 2011. These observational measurements within the photic zone demonstrated the simultaneous regeneration and assimilation of NH4+, NO2− and NO3−. NH4+ was assimilated at 1.82–49.12 nmol N L−1 h−1 and regenerated at 3.46–14.60 nmol N L−1 h−1; NO2− was assimilated at 0–2.08 nmol N L−1 h−1 and regenerated at 0.01–1.85 nmol N L−1 h−1; NO3− was assimilated at 0.67–18.75 nmol N L−1 h−1 and regenerated at 0.05–28.97 nmol N L−1 h−1. Observations implied that these processes were closely coupled at the regional scale and nitrogen recycling played an important role in sustaining phytoplankton growth during the summer. The [N2O], measured in water column profiles, was 10.13 ± 1.11 nmol L−1 and did not strongly diverge from atmospheric equilibrium indicating that sampled marine regions where neither a strong source nor sink of N2O to the atmosphere. Multivariate analysis of data describing water column biogeochemistry and its links to N-cycling activity failed to explain the observed variance in rates of N-regeneration and N-assimilation, possibly due to the limited number of process rate observations. In the surface waters of 5 further stations, Ocean Acidification (OA) bioassay experiments were conducted to investigate the response of NH4+ oxidising and regenerating organisms to simulated OA conditions, including the implications for [N2O]. Multivariate analysis was undertaken which considered the complete bioassay dataset of measured variables describing changes in N-regeneration rate, [N2O] and the biogeochemical composition of seawater. While anticipating biogeochemical differences between locations, we aimed to test the hypothesis that the underlying mechanism through which pelagic N-regeneration responded to simulated OA conditions was independent of location and that a mechanistic understanding of how NH4+ oxidation, NH4+ regeneration and N2O production responded to OA could be developed. Results indicated that N-regeneration process responses to OA treatments were location specific; no mechanistic understanding of how N-regeneration processes respond to OA in the surface ocean of the NW European shelf sea could be developed.

Biomass changes and trophic amplification of plankton in a warmer ocean

Ocean warming can modify the ecophysiology and distribution of marine organisms, and relationships between species, with nonlinear interactions between ecosystem components potentially resulting in trophic amplification. Trophic amplification (or attenuation) describe the propagation of a hydroclimatic signal up the food web, causing magnification (or depression) of biomass values along one or more trophic pathways. We have employed 3-D coupled physical-biogeochemical models to explore ecosystem responses to climate change with a focus on trophic amplification. The response of phytoplankton and zooplankton to global climate-change projections, carried out with the IPSL Earth System Model by the end of the century, is analysed at global and regional basis, including European seas (NE Atlantic, Barents Sea, Baltic Sea, Black Sea, Bay of Biscay, Adriatic Sea, Aegean Sea) and the Eastern Boundary Upwelling System (Benguela). Results indicate that globally and in Atlantic Margin and North Sea, increased ocean stratification causes primary production and zooplankton biomass to decrease in response to a warming climate, whilst in the Barents, Baltic and Black Seas, primary production and zooplankton biomass increase. Projected warming characterized by an increase in sea surface temperature of 2.29 ± 0.05 °C leads to a reduction in zooplankton and phytoplankton biomasses of 11% and 6%, respectively. This suggests negative amplification of climate driven modifications of trophic level biomass through bottom-up control, leading to a reduced capacity of oceans to regulate climate through the biological carbon pump. Simulations suggest negative amplification is the dominant response across 47% of the ocean surface and prevails in the tropical oceans; whilst positive trophic amplification prevails in the Arctic and Antarctic oceans. Trophic attenuation is projected in temperate seas. Uncertainties in ocean plankton projections, associated to the use of single global and regional models, imply the need for caution when extending these considerations into higher trophic levels.

The dispersal of phytoplankton populations by enhanced turbulent mixing in a shallow coastal sea

A single tidal cycle survey in a Lagrangian reference frame was conducted in autumn 2010 to evaluate the impact of short-term, episodic and enhanced turbulent mixing on large chain-forming phytoplankton. Observations of turbulence using a free-falling microstructure profiler were undertaken, along with near-simultaneous profiles with an in-line digital holographic camera at station L4 (50° 15′ N 4° 13′ W, depth 50 m) in the Western English Channel. Profiles from each instrument were collected hourly whilst following a drogued drifter. Results from an ADCP attached to the drifter showed pronounced vertical shear, indicating that the water column structure consisted of two layers, restricting interpretation of the Lagrangian experiment to the upper ~ 25 m. Atmospheric conditions deteriorated during the mid-point of the survey, resulting in values of turbulent dissipation reaching a maximum of 10− 4 W kg− 1 toward the surface in the upper 10 m. Chain-forming phytoplankton > 200 μm were counted using the data from the holographic camera for the two periods, before and after the enhanced mixing event. As mixing increased phytoplankton underwent chain breakage, were dispersed by advection through their removal from the upper to lower layer and subjected to aggregation with other suspended material. Depth averaged counts of phytoplankton were reduced from a maximum of around 2050 L− 1 before the increased turbulence, to 1070 L− 1 after, with each of these mechanisms contributing to this reduction. These results demonstrate the sensitivity of phytoplantkon populations to moderate increases in turbulent activity, yielding consequences for accurate forecasting of the role played by phytoplankton in climate studies and also for the ecosystem in general in their role as primary producers.

The influence of ocean acidification on nitrogen regeneration and nitrous oxide production in the North-West European shelf sea

The assimilation and regeneration of dissolved inorganic nitrogen, and the concentration of N2O, was investigated at stations located in the NW European shelf sea during June/July 2011. These observational measurements within the photic zone demonstrated the simultaneous regeneration and assimilation of NH4+, NO2− and NO3−. NH4+ was assimilated at 1.82–49.12 nmol N L−1 h−1 and regenerated at 3.46–14.60 nmol N L−1 h−1; NO2− was assimilated at 0–2.08 nmol N L−1 h−1 and regenerated at 0.01–1.85 nmol N L−1 h−1; NO3− was assimilated at 0.67–18.75 nmol N L−1 h−1 and regenerated at 0.05–28.97 nmol N L−1 h−1. Observations implied that these processes were closely coupled at the regional scale and nitrogen recycling played an important role in sustaining phytoplankton growth during the summer. The [N2O], measured in water column profiles, was 10.13 ± 1.11 nmol L−1 and did not strongly diverge from atmospheric equilibrium indicating that sampled marine regions where neither a strong source nor sink of N2O to the atmosphere. Multivariate analysis of data describing water column biogeochemistry and its links to N-cycling activity failed to explain the observed variance in rates of N-regeneration and N-assimilation, possibly due to the limited number of process rate observations. In the surface waters of 5 further stations, Ocean Acidification (OA) bioassay experiments were conducted to investigate the response of NH4+ oxidising and regenerating organisms to simulated OA conditions, including the implications for [N2O]. Multivariate analysis was undertaken which considered the complete bioassay dataset of measured variables describing changes in N-regeneration rate, [N2O] and the biogeochemical composition of seawater. While anticipating biogeochemical differences between locations, we aimed to test the hypothesis that the underlying mechanism through which pelagic N-regeneration responded to simulated OA conditions was independent of location and that a mechanistic understanding of how NH4+ oxidation, NH4+ regeneration and N2O production responded to OA could be developed. Results indicated that N-regeneration process responses to OA treatments were location specific; no mechanistic understanding of how N-regeneration processes respond to OA in the surface ocean of the NW European shelf sea could be developed.

The influence of ocean acidification on nitrogen regeneration and nitrous oxide production in the northwest European shelf sea

The assimilation and regeneration of dissolved inorganic nitrogen, and the concentration of N₂O, was investigated at stations located in the NW European shelf sea during June/July 2011. These observational measurements within the photic zone demonstrated the simultaneous regeneration and assimilation of NH₄⁺, NO₂⁻ and NO₃⁻. NH₄⁺ was assimilated at 1.82–49.12 nmol N L⁻¹ h⁻¹ and regenerated at 3.46–14.60 nmol N L⁻¹ h⁻¹; NO₂^- was assimilated at 0–2.08 nmol N L⁻¹ h⁻¹ and regenerated at 0.01–1.85 nmol N L⁻¹ h⁻¹; NO₃⁻ was assimilated at 0.67–18.75 nmol N L⁻¹ h⁻¹ and regenerated at 0.05–28.97 nmol N L⁻¹ h⁻¹. Observations implied that these processes were closely coupled at the regional scale and that nitrogen recycling played an important role in sustaining phytoplankton growth during the summer. The [N₂O], measured in water column profiles, was 10.13 ± 1.11 nmol L⁻¹ and did not strongly diverge from atmospheric equilibrium indicating that sampled marine regions were neither a strong source nor sink of N₂O to the atmosphere. Multivariate analysis of data describing water column biogeochemistry and its links to N-cycling activity failed to explain the observed variance in rates of N-regeneration and N-assimilation, possibly due to the limited number of process rate observations. In the surface waters of five further stations, ocean acidification (OA) bioassay experiments were conducted to investigate the response of NH₄⁺ oxidising and regenerating organisms to simulated OA conditions, including the implications for [N₂O]. Multivariate analysis was undertaken which considered the complete bioassay data set of measured variables describing changes in N-regeneration rate, [N₂O] and the biogeochemical composition of seawater. While anticipating biogeochemical differences between locations, we aimed to test the hypothesis that the underlying mechanism through which pelagic N-regeneration responded to simulated OA conditions was independent of location. Our objective was to develop a mechanistic understanding of how NH₄⁺ regeneration, NH₄⁺ oxidation and N₂O production responded to OA. Results indicated that N-regeneration process responses to OA treatments were location specific; no mechanistic understanding of how N-regeneration processes respond to OA in the surface ocean of the NW European shelf sea could be developed.

Factors governing the deep ventilation of the Red Sea

A variety of data based on hydrographic measurements, satellite observations, reanalysis databases, and meteorological observations are used to explore the interannual variability and factors governing the deep water formation in the northern Red Sea. Historical and recent hydrographic data consistently indicate that the ventilation of the near-bottom layer in the Red Sea is a robust feature of the thermohaline circulation. Dense water capable to reach the bottom layers of the Red Sea can be regularly produced mostly inside the Gulfs of Aqaba and Suez. Occasionally, during colder than usual winters, deep water formation may also take place over coastal areas in the northernmost end of the open Red Sea just outside the Gulfs of Aqaba and Suez. However, the origin as well as the amount of deep waters exhibit considerable interannual variability depending not only on atmospheric forcing but also on the water circulation over the northern Red Sea. Analysis of several recent winters shows that the strength of the cyclonic gyre prevailing in the northernmost part of the basin can effectively influence the sea surface temperature (SST) and intensify or moderate the winter surface cooling. Upwelling associated with periods of persistent gyre circulation lowers the SST over the northernmost part of the Red Sea and can produce colder than normal winter SST even without extreme heat loss by the sea surface. In addition, the occasional persistence of the cyclonic gyre feeds the surface layers of the northern Red Sea with nutrients, considerably increasing the phytoplankton biomass.