Metabolically active, non-nitrogen fixing,Trichodesmiumin UK coastal waters during winter

Trichodesmium, a colonial cyanobacterium typically associated with tropical waters, was observed between January and April 2014 in the western English Channel. Sequencing of the heterocyst differentiation (hetR) and 16S rRNA genes placed this community within the Clade IV Trichodesmium, an understudied clade previously found only in low numbers in warmer waters. Nitrogen fixation was not detected although measurable rates of nitrate uptake and carbon fixation were observed. Trichodesmium RuBisCO transcript abundance relative to gene abundance suggests the potential for viable and potentially active Trichodesmium carbon fixation. Observations of Trichodesmium when coupled with a numerical advection model indicate that Trichodesmium communities can remain viable for >3.5 months at temperatures lower than previously expected. The results suggest that Clade IV Trichodesmium occupies a different niche to other Trichodesmium species, and is a cold- or low-light-adapted variant.

Potential impacts of climate change on the primary production of regional seas: A comparative analysis of five European seas

Regional seas are potentially highly vulnerable to climate change, yet are the most directly societally important regions of the marine environment. The combination of widely varying conditions of mixing, forcing, geography (coastline and bathymetry) and exposure to the open-ocean makes these seas subject to a wide range of physical processes that mediates how large scale climate change impacts on these seas’ ecosystems. In this paper we explore the response of five regional sea areas to potential future climate change, acting via atmospheric, oceanic and terrestrial vectors. These include the Barents Sea, Black Sea, Baltic Sea, North Sea, Celtic Seas, and are contrasted with a region of the Northeast Atlantic. Our aim is to elucidate the controlling dynamical processes and how these vary between and within these seas. We focus on primary production and consider the potential climatic impacts on: long term changes in elemental budgets, seasonal and mesoscale processes that control phytoplankton’s exposure to light and nutrients, and briefly direct temperature response. We draw examples from the MEECE FP7 project and five regional model systems each using a common global Earth System Model as forcing. We consider a common analysis approach, and additional sensitivity experiments. Comparing projections for the end of the 21st century with mean present day conditions, these simulations generally show an increase in seasonal and permanent stratification (where present). However, the first order (low- and mid-latitude) effect in the open ocean projections of increased permanent stratification leading to reduced nutrient levels, and so to reduced primary production, is largely absent, except in the NE Atlantic. Even in the two highly stratified, deep water seas we consider (Black and Baltic Seas) the increase in stratification is not seen as a first order control on primary production. Instead, results show a highly heterogeneous picture of positive and negative change arising from complex combinations of multiple physical drivers, including changes in mixing, circulation and temperature, which act both locally and non-locally through advection.

Monsoon oscillations regulate fertility of the Red Sea

Tropical ocean ecosystems are predicted to become warmer, more saline, and less fertile in a future Earth. The Red Sea, one of the warmest and most saline environments in the world, may afford insights into the function of the tropical ocean ecosystem in a changing planet. We show that the concentration of chlorophyll and the duration of the phytoplankton growing season in the Red Sea are controlled by the strength of the winter Arabian monsoon (through horizontal advection of fertile waters from the Indian Ocean). Furthermore, and contrary to expectation, in the last decade (1998–2010) the winter Red Sea phytoplankton biomass has increased by 75% during prolonged positive phases of the Multivariate El Niño–Southern Oscillation Index. A new mechanism is reported, revealing the synergy of monsoon and climate in regulating Red Sea greenness.

The significance of nitrogen regeneration for new production within a filament of the Mauritanian upwelling system

The Lagrangian progression of a biological community was followed in a filament of the Mauritanian upwelling system, north-west Africa, during offshore advection. The inert dual tracers sulfur hexafluoride and helium-3 labelled a freshly upwelled patch of water that was mapped for 8 days. Changes in biological, physical, and chemical characteristics were measured, including phytoplankton productivity, nitrogen assimilation, and regeneration. Freshly upwelled water contained high nutrient concentrations but was depleted in N compared to Redfield stoichiometry. The highest rate of primary productivity was measured on the continental shelf, associated with high rates of nitrogen assimilation and a phytoplankton community dominated by diatoms and flagellates. Indicators of phytoplankton abundance and activity decreased as the labelled water mass transited the continental shelf slope into deeper water, possibly linked to the mixed layer depth exceeding the light penetration depth. By the end of the study, the primary productivity rate decreased and was associated with lower rates of nitrogen assimilation and lower nutrient concentrations. Nitrogen regeneration and assimilation took place simultaneously. Results highlighted the importance of regenerated NHC 4 in sustaining phytoplankton productivity and indicate that the upwelled NO3 pool contained an increasing fraction of regenerated NO3 as it advected offshore. By calculating this fraction and incorporating it into an f ratio formulation, we estimated that of the 12:38Tg C of annual regional production, 4:73Tg C was exportable.

The significance of nitrogen regeneration for new production within a filament of the Mauritanian upwelling system

The Lagrangian progression of a biological community was followed in a filament of the Mauritanian upwelling system, north-west Africa, during offshore advection. The inert dual tracers sulfur hexafluoride and helium-3 labelled a freshly upwelled patch of water that was mapped for 8 days. Changes in biological, physical, and chemical characteristics were measured, including phytoplankton productivity, nitrogen assimilation, and regeneration. Freshly upwelled water contained high nutrient concentrations but was depleted in N compared to Redfield stoichiometry. The highest rate of primary productivity was measured on the continental shelf, associated with high rates of nitrogen assimilation and a phytoplankton community dominated by diatoms and flagellates. Indicators of phytoplankton abundance and activity decreased as the labelled water mass transited the continental shelf slope into deeper water, possibly linked to the mixed layer depth exceeding the light penetration depth. By the end of the study, the primary productivity rate decreased and was associated with lower rates of nitrogen assimilation and lower nutrient concentrations. Nitrogen regeneration and assimilation took place simultaneously. Results highlighted the importance of regenerated NHC 4 in sustaining phytoplankton productivity and indicate that the upwelled NO3 pool contained an increasing fraction of regenerated NO3 as it advected offshore. By calculating this fraction and incorporating it into an f ratio formulation, we estimated that of the 12:38Tg C of annual regional production, 4:73Tg C was exportable.

Individual-based modelling of the development and transport of a Karenia mikimotoi bloom on the North-West European continental shelf

In 2006, a large and prolonged bloom of the dinoflagellate Karenia mikimotoi occurred in Scottish coastal waters, causing extensive mortalities of benthic organisms including annelids and molluscs and some species of fish ( Davidson et al., 2009). A coupled hydrodynamic-algal transport model was developed to track the progression of the bloom around the Scottish coast during June–September 2006 and hence investigate the processes controlling the bloom dynamics. Within this individual-based model, cells were capable of growth, mortality and phototaxis and were transported by physical processes of advection and turbulent diffusion, using current velocities extracted from operational simulations of the MRCS ocean circulation model of the North-west European continental shelf. Vertical and horizontal turbulent diffusion of cells are treated using a random walk approach. Comparison of model output with remotely sensed chlorophyll concentrations and cell counts from coastal monitoring stations indicated that it was necessary to include multiple spatially distinct seed populations of K. mikimotoi at separate locations on the shelf edge to capture the qualitative pattern of bloom transport and development. We interpret this as indicating that the source population was being transported northwards by the Hebridean slope current from where colonies of K. mikimotoi were injected onto the continental shelf by eddies or other transient exchange processes. The model was used to investigate the effects on simulated K. mikimotoi transport and dispersal of: (1) the distribution of the initial seed population; (2) algal growth and mortality; (3) water temperature; (4) the vertical movement of particles by diurnal migration and eddy diffusion; (5) the relative role of the shelf edge and coastal currents; (6) the role of wind forcing. The numerical experiments emphasized the requirement for a physiologically based biological model and indicated that improved modelling of future blooms will potentially benefit from better parameterisation of temperature dependence of both growth and mortality and finer spatial and temporal hydrodynamic resolution.

Individual-based modelling of the development and transport of a Karenia mikimotoi bloom on the North-West European continental shelf

In 2006, a large and prolonged bloom of the dinoflagellate Karenia mikimotoi occurred in Scottish coastal waters, causing extensive mortalities of benthic organisms including annelids and molluscs and some species of fish ( Davidson et al., 2009). A coupled hydrodynamic-algal transport model was developed to track the progression of the bloom around the Scottish coast during June–September 2006 and hence investigate the processes controlling the bloom dynamics. Within this individual-based model, cells were capable of growth, mortality and phototaxis and were transported by physical processes of advection and turbulent diffusion, using current velocities extracted from operational simulations of the MRCS ocean circulation model of the North-west European continental shelf. Vertical and horizontal turbulent diffusion of cells are treated using a random walk approach. Comparison of model output with remotely sensed chlorophyll concentrations and cell counts from coastal monitoring stations indicated that it was necessary to include multiple spatially distinct seed populations of K. mikimotoi at separate locations on the shelf edge to capture the qualitative pattern of bloom transport and development. We interpret this as indicating that the source population was being transported northwards by the Hebridean slope current from where colonies of K. mikimotoi were injected onto the continental shelf by eddies or other transient exchange processes. The model was used to investigate the effects on simulated K. mikimotoi transport and dispersal of: (1) the distribution of the initial seed population; (2) algal growth and mortality; (3) water temperature; (4) the vertical movement of particles by diurnal migration and eddy diffusion; (5) the relative role of the shelf edge and coastal currents; (6) the role of wind forcing. The numerical experiments emphasized the requirement for a physiologically based biological model and indicated that improved modelling of future blooms will potentially benefit from better parameterisation of temperature dependence of both growth and mortality and finer spatial and temporal hydrodynamic resolution.

Surface water subduction during a downwelling event in a semi-enclosed bay

The Ría de Vigo is a bay strongly influenced by upwelling-downwelling cycles along the adjacent coast of NW Iberia. Moored and ship-board observations during September 2006 showed that subduction, initially associated with an estuarine circulation, strengthened when a strong downwelling circulation, resulting from northward wind over the coastal ocean, was generated in the outer ría causing ambient waters to be advected outward in the lower layer. Incoming surface waters confined the estuarine circulation to the shallow interior and displaced isopleths downward through the water column at ∼10 m d−1. As the estuarine circulation retreated inward, strong flow convergence developed between middle and inner ria in the layer above 15 m, while divergence developed beneath. The convergence increased through the period of downwelling-favorable wind at a rate consistent with the observed isopleth displacement velocities. The coefficient of turbulent diffusion Kt, from a microstructure profiler, indicated that mixing was strong in the estuarine circulation and subsequently in the downwelling zone, where localized instabilities and temperature-salinity inversions were observed. During the downwelling, concentrations of phytoplankton, including potentially harmful species, increased, especially in the middle and inner ria, as a result of inward advection, subduction and the ability of the dinoflagellates to maintain their position in the water column by swimming. In the course of the 5 day event, the water mass of all but the innermost ría was flushed completely and replaced by waters originating in the coastally-trapped poleward flow along the Atlantic coastline.

Surface water subduction during a downwelling event in a semi-enclosed bay

The Ría de Vigo is a bay strongly influenced by upwelling-downwelling cycles along the adjacent coast of NW Iberia. Moored and ship-board observations during September 2006 showed that subduction, initially associated with an estuarine circulation, strengthened when a strong downwelling circulation, resulting from northward wind over the coastal ocean, was generated in the outer ría causing ambient waters to be advected outward in the lower layer. Incoming surface waters confined the estuarine circulation to the shallow interior and displaced isopleths downward through the water column at ∼10 m d−1. As the estuarine circulation retreated inward, strong flow convergence developed between middle and inner ria in the layer above 15 m, while divergence developed beneath. The convergence increased through the period of downwelling-favorable wind at a rate consistent with the observed isopleth displacement velocities. The coefficient of turbulent diffusion Kt, from a microstructure profiler, indicated that mixing was strong in the estuarine circulation and subsequently in the downwelling zone, where localized instabilities and temperature-salinity inversions were observed. During the downwelling, concentrations of phytoplankton, including potentially harmful species, increased, especially in the middle and inner ria, as a result of inward advection, subduction and the ability of the dinoflagellates to maintain their position in the water column by swimming. In the course of the 5 day event, the water mass of all but the innermost ría was flushed completely and replaced by waters originating in the coastally-trapped poleward flow along the Atlantic coastline.