Description of a flexible and extendable physical–biogeochemical model system for the water column

A modelling system for coupled physical–biogeochemical simulations in the water column is presented here. The physical model component allows for a number of different statistical turbulence closure schemes, ranging from simple algebraic closures to two-equation turbulence models with algebraic second-moment closures. The biogeochemical module consists of models which are based on a number of state variables represented by their ensemble averaged concentrations. Specific biogeochemical models may range from simple NPZ (nutrient–phytoplankton–zooplankton) to complex ecosystem models. Recently developed modified Patankar solvers for ordinary differential equations allow for stable discretisations of the production and destruction terms guaranteeing conservative and non-negative solutions. The increased stability of these new solvers over explicit solvers is demonstrated for a plankton spring bloom simulation. The model system is applied to marine ecosystem dynamics the Northern North Sea and the Central Gotland Sea. Two different biogeochemical models are applied, a conservative nitrogen-based model to the North Sea, and a more complex model including an oxygen equation to the Baltic Sea, allowing for the reproduction of chemical processes under anoxic conditions. For both applications, earlier model results obtained with slightly different model setups could be basically reproduced. It became however clear that the choice for ecosystem model parameters such as maximum phytoplankton growth rates does strongly depend on the physical model parameters (such as turbulence closure models or external forcing).     

Error quantification of a high resolution coupled hydrodynamic-ecosystem coastal-ocean model: Part3, validation with Continuous Plankton Recorder data

Data collected during the Continuous Plankton Recorder (CPR) survey has been used to validate a three-dimensional hydrodynamic ecosystem model simulation of the North-west European Shelf for the years 1988–89. The CPR time series is unique to the North Atlantic region as a validation tool. Data were extracted from the model to correspond with those collected by the CPR survey, and both the model and survey plankton data were standardised to allow the comparison of model biomass with survey counts. Simple linear regression and absolute error maps provide a qualitative evaluation of spatio-temporal model performance of simulated diatoms, flagellates, total phytoplankton and omnivorous mesozooplankton. Comparisons of z-scores indicate that the model reproduces the main pelagic seasonal features, and there is good correlation between magnitudes of these features with respect to standard deviations from a long-term mean. The model is replicating up to 62% of the mesozooplankton seasonality across the domain, with variable results for the phytoplankton. There are, however, differences in the timing of patterns in plankton seasonality. The validation exercise has highlighted that the spring diatom bloom in the model is too early, suggesting the need to reparameterise the response of phytoplankton to changing light levels in the model. Errors in the north and west of the domain imply that model turbulence and vertical density structure need to be improved to more accurately capture plankton dynamics.     

One-dimensional modelling of the plankton ecosystem of the north-western Corsican coastal area in relation to meteorological constraints

In order to study the influence of wind mixing on the spring variability of the plankton production of the north western Corsican coastal area, a one-dimensional (1D), vertical, coupled hydrodynamic/biological model (ECOHYDROMV) is used. A hydrodynamic 1D model of the water column with a k–l turbulent closure is applied. The biological model comprises six state variables, representing the plankton ecosystem in the spring period: phytoplankton, copepods, nitrate, ammonium, particulate organic matter of phytoplanktonic origin and particulate organic matter of zooplanktonic origin. The system is influenced by turbulence (expressed by the vertical eddy diffusivity), temperature and irradiance. The model takes into account momentum and heat surface fluxes computed from meteorological data in order to simulate a typical spring atmospheric forcing for the considered area. Results show that primary production vertical structure is characterised by a subsurface maximum which deepens with time and is regulated by the opposite gradients of nitrate concentration and irradiance. Surface plankton productivity is mainly controlled by turbulent vertical transport of nutrients into the mixed layer. The short time scale variability of turbulent mixing generated by the wind appears to be responsible for the plurimodal shape of plankton blooms, observed in the considered area. Furthermore, the model is applied to the study of the spring evolution of the plankton communities off the bay of Calvi (Corsica) for the years 1986 and 1988. In order to initiate and validate the model, time series of hydrological, chemical and biological data have been used. The model reproduces accurately the spring evolution of the phytoplankton biomass measured in situ and illustrates that its strong variability in those years was in close relation to the variability of the wind intensity.     

Application of new parameterizations of gas transfer velocity and their impact on regional and global marine CO2 budgets

One of the dominant sources of uncertainty in the calculation of air–sea flux of carbon dioxide on a global scale originates from the various parameterizations of the gas transfer velocity, k, that are in use. Whilst it is undisputed that most of these parameterizations have shortcomings and neglect processes which influence air–sea gas exchange and do not scale with wind speed alone, there is no general agreement about their relative accuracy.The most widely used parameterizations are based on non-linear functions of wind speed and, to a lesser extent, on sea surface temperature and salinity. Processes such as surface film damping and whitecapping are known to have an effect on air–sea exchange. More recently published parameterizations use friction velocity, sea surface roughness, and significant wave height. These new parameters can account to some extent for processes such as film damping and whitecapping and could potentially explain the spread of wind-speed based transfer velocities published in the literature.We combine some of the principles of two recently published k parameterizations [Glover, D.M., Frew, N.M., McCue, S.J. and Bock, E.J., 2002. A multiyear time series of global gas transfer velocity from the TOPEX dual frequency, normalized radar backscatter algorithm. In: Donelan, M.A., Drennan, W.M., Saltzman, E.S., and Wanninkhof, R. (Eds.), Gas Transfer at Water Surfaces, Geophys. Monograph 127. AGU,Washington, DC, 325–331; Woolf, D.K., 2005. Parameterization of gas transfer velocities and sea-state dependent wave breaking. Tellus, 57B: 87–94] to calculate k as the sum of a linear function of total mean square slope of the sea surface and a wave breaking parameter. This separates contributions from direct and bubble-mediated gas transfer as suggested by Woolf [Woolf, D.K., 2005. Parameterization of gas transfer velocities and sea-state dependent wave breaking. Tellus, 57B: 87–94] and allows us to quantify contributions from these two processes independently.We then apply our parameterization to a monthly TOPEX altimeter gridded 1.5° × 1.5° data set and compare our results to transfer velocities calculated using the popular wind-based k parameterizations by Wanninkhof [Wanninkhof, R., 1992. Relationship between wind speed and gas exchange over the ocean. J. Geophys. Res., 97: 7373–7382.] and Wanninkhof and McGillis [Wanninkhof, R. and McGillis, W., 1999. A cubic relationship between air−sea CO2 exchange and wind speed. Geophys. Res. Lett., 26(13): 1889–1892]. We show that despite good agreement of the globally averaged transfer velocities, global and regional fluxes differ by up to 100%. These discrepancies are a result of different spatio-temporal distributions of the processes involved in the parameterizations of k, indicating the importance of wave field parameters and a need for further validation.     

Physical–biological coupling in spore dispersal of kelp forest macroalgae

The physical–biological linkages controlling the dispersal of spores produced by macroalgae that reside in kelp forests are complicated and laced with feedbacks. Here we discuss the fundamental elements of these interactions. Biological considerations include spore swimming and sinking speeds, their periods of viability in the plankton, and the height of spore release above the seafloor, which together determine the durations over which spores can be swept by horizontal currents before they contact the seafloor. Morphologies and material properties of canopy forming kelps may also influence the drag exerted on passing waters by the kelps, the plants' ability to persist in the face of rapid flows, and thereby the degree to which impinging currents are redirected around, or slowed within, kelp forests. Macroalgal life histories, and the size of spore sources as controlled by the dimensions of kelp forests and the density and fecundity of individuals within them, influence effective dispersal distances as well. Physical considerations encompass the mean speed, direction, and timescales of variability of currents relative to spore suspension times, the interaction of surface gravity waves with currents in producing turbulence in the benthic boundary layer, wind-driven surface mixing, water stratification, and shoreline bathymetry and substratum roughness, all of which can affect the interplay of vertical and horizontal transport of macroalgal spores. Intricate within-forest processes may induce attenuation of current speeds and consequent reductions in seabed shear, along with simultaneous production of small-scale turbulence in kelp wakes. Slower mean currents and smaller eddy scales in turn may attenuate vertical mixing within forests, thus extending spore suspension times. Further complexities likely arise due to changes in the relative rates of horizontal and vertical dispersion, modifications to the overall profiles of vertical mixing, and the creation of fine-scale secondary flows around kelp individuals and substratum features. Under conditions of more rapid currents, submergence of the surface canopy and the establishment of skimming flows at the canopy–fluid interface may introduce additional coherent flow structures that alter rates of fluid exchange to and from the forest. Many of these coupled physical–biological processes are just beginning to be examined in a rigorous fashion in kelp forests, but their potential importance is clear.     

The influence of different turbulence schemes on modelling primary production in a 1D coupled physical–biological model

A one-dimensional (1D) coupled physical–microbiological model has been applied to a site in the central North Sea. The impact of the choice of the turbulence closure scheme on the modelling the primary production has been investigated.The model was run with four different parameterisations of vertical mixing of heat, momentum and dissolved and suspended matters, using M2 tidal forcing and the hourly mean meteorological forcing of 1989 to reproduce the annual thermal structure and primary production. The four mixing parameterisations are: Level 2 turbulence closure scheme [Mellor, G.L., Yamada, T., 1974. A hierarchy of turbulence closure models for planetary boundary layers. J. Atmos. Sci. 31, 1791–1806; Mellor, G.L., Yamada, T., 1982. Development of a turbulence closure model for geophysical Fluid problems. Rev. Geophys. Space Phys. 20 (4) 851–875] using an explicit numerical scheme [Sharples, J., Tett, P., 1994. Modelling the effect of physical variability on the midwater chlorophyll maximum. J. Mar. Res. 52, 219–238]; a version of the Level 2.5 turbulence closure scheme [Galperin, B., Kantha, L.H., Hassid, S., Rosati, A., 1988. A quasi-equilibrium turbulent energy model for geophysical flows. J. Atmos. Sci. 45, 55–62; Ruddick, K.G., Deleersnijder, E., Luyten, P.J., Ozer, J., 1995. Haline stratification in the rhine/meuse freshwater plume: a 3D model sensitivity analysis. Cont. Shelf Res. 15 (13) 1597–1630] simplified to use an algebraic mixing length by Sharples and Simpson [Sharples, J., Simpson, J.H., 1995. Semidiurnal and longer period stability cycles in the Liverpool Bay region of freshwater influence. Cont. Shelf Res. 15, 295–313], also solved explicitly; the same simplified L2.5 scheme with an implicit numerical solution and modified vertical discretisation scheme [Annan, J.D., 1999. Numerical methods for the solution of the turbulence energy equations in the shelf seas. Int. J. Numer. Methods Fluids 29, 193–206]; and another version of the same scheme (but using a different algebraic mixing length) as described by Xing and Davies [Xing, J., Davies, A.M., 1996a. Application of turbulence energy models to the computation of tidal currents and mixing intensities in the shelf edge regions. J. Phys. Oceanogr. 26, 417–447; Xing, J., Davies, A.M., 1996b. Application of a range of turbulence models to the computation of tidal currents and mixing intensities in shelf edge regions. Cont. Shelf. Res. 16, 517–547; Xing, J., Davies, A.M., 1998. Application of a range of turbulence energy models to the computation of the internal tide. Int. J. Numer. Methods Fluids 26, 1055–1084]. Various model outputs at the sea surface and in depth profiles have been compared with data collected in 1989 as part of the North Sea Project [Huthnance, J.M., 1990. Progress on North Sea Project. NERC News, vol. 12, pp. 25–29, UK]. It is shown that the biological results are extremely sensitive to the small changes in the physical conditions, which arise due to the different turbulence schemes tested. The timing of the spring bloom and the maintenance of the midwater chlorophyll maximum all differ greatly between model runs, and the gross primary production varies by a factor of two from the highest to lowest results. The simplified Level 2.5 scheme, implemented using the numerical methods of Annan [Annan, J.D., 1999. Numerical methods for the solution of the turbulence energy equations in the shelf seas. Int. J. Numer. Methods Fluids 29, 193–206], produces results, which give the best agreement with the available data.     

Intermittent turbulence and copepod dynamics: Increase in encounter rates through preferential concentration

Turbulence has a strong influence on plankton contact rate, which is a crucial parameter for plankton ecology. In the field of particle-turbulence interactions, it is now well known that fully developed turbulence does not always homogenise particle distributions, but instead creates, in some well-defined conditions, preferential concentrations of heavy particles. Many studies have considered the influence of this type of preferential concentration on particle contact rate. We consider here the possible application of these results to copepods, assuming that some results obtained for heavy particles are still valid for light particles. Using parameter values representative of copepod species in coastal waters and open ocean, we numerically estimate the possible enhancement of copepod contact rates due to the preferential concentration by turbulence. The assessment is done by using data from a laboratory experiment: we find from the trajectory analysis of small neutrally buoyant particles, that the preferential concentration effect increases the contact rate up to 140%. We argue that this effect may be more pronounced for higher Reynolds numbers, and may have important ecological applications.     

Quantitative estimation of the influence of surface thermal fronts over chlorophyll concentration at the Patagonian shelf

Eighteen-year (1985–2002) mean monthly SST Pathfinder data with 9 km spatial resolution have been used to estimate surface gradients by finite differences. Then the seasonal climatological means have been calculated from the intensity of these gradients, and surface thermal fronts present in the Patagonian Continental Shelf (PCS) have been located. Moreover, 6 years (1998–2003) of SeaWiFS data with approximately 4 km spatial resolution have been used to estimate monthly composite images of surface chlorophyll concentration, after which seasonal climatological means distributions have been generated. Both seasonal distributions have been analyzed together and by combining the knowledge of oceanographic processes and phytoplankton responses to light and nutrient availability, regions where the presence of a thermal front affects photosynthetic activity have been identified. Subjective criteria have been applied to define eighteen areas where phytoplankton biomass is influenced by the presence of a thermal front. In these areas, the surface chlorophyll (spatial mean and total), its relationship with the surface chlorophyll of the whole region, and the seasonal evolution of this relationship have been calculated. All frontal areas cover less than 15% of the total surface, but they contribute with over 23% of the phytoplankton annual mean biomass. Considered as a group, during summer they show high chlorophyll values very similar to those in spring. During the cold period, when the water column is vertically mixed in practically the whole of PCS, the influence of physical fronts over the biological production is minimum. The frontal zone image remains clearly defined during summer, when approximately 85% of the area will have a determined mean chlorophyll concentration, while the other 15% has a 2.45 times larger value. While three pattern trends have been identified in the frontal areas, only two of them condition the pattern of the group, due to their horizontal extension.     

The importance of oceanographic fronts to marine birds and mammals of the southern oceans

During the last 30 years, at-sea studies of seabirds and marine mammals in the oceans south of the Subtropical Front have described an association with major frontal areas. More recently, the advancement in microtechnology has allowed the tracking of individuals and investigations into how these marine predators actually use the frontal zones. In this review, we examine 1) the relative importance to apex predators of the different frontal zones in terms of spatial distribution and carbon flux; 2) the processes that determine their preferential use; and 3) how the mesoscale dynamics of frontal structures drive at-sea foraging strategies of these predators. We review published results from southern waters and place them in a broader context with respect to what has been learned about the importance of fronts in oceans farther north.Some fronts constitute important boundaries for seabird communities in southern waters. At a mesoscale the maximum values of seabird diversity and abundance correspond to the location of the main fronts. At-sea surveys show a strong curvilinear correlation between seabird abundance and sea surface temperatures. High mean species richness and diversity for whales and seabirds are consistently associated with the southern water mass boundary of the Antarctic Circumpolar Current, the Subtropical Front and the Subantarctic Front; in the case of the Polar Front mean seabird densities are more variable. At small-scales, variation in seabird occurrence has been directly related to the processes at fronts in a limited number of cases. A significant positive relation was found between some plankton feeding species and frontal temperature gradient–phytoplankton variables.Telemetric studies have revealed that several apex predators (penguins, albatrosses, seals) perform long, directed foraging trips either to the Subtropical front or Polar Front, depending on locality. Seabirds with low flight costs, such as albatrosses, are able to reach fronts at long distances from colonies, showing variable foraging strategies as a function of the distances involved. Diving birds such as King penguins, that travel at a higher cost and lower speed, rely on the predictable spatial distribution of mesopelagic fish found close to the Polar Front. They may use the currents associated with eddies as oceanographic cues in the active search for frontal zones. Once in these areas they dive preferentially in and below the depth of the thermocline where catches per unit effort are high. Elephant seals concentrate foraging activity principally inside or at the boundary of cyclonic eddies. These mesoscale features appear to offer exceptional productivity favourable for foraging by various diving top predators.The connection between biophysical parameters at fronts and predators is likely to be made through biological enhancement. Top predators appear to forage at locations where prey are advected by physical processes and others where prey are produced locally. Long-term research on at-sea distributions and demographic parameters of top predators are essential to assess the consequences of potential shift in front distributions in relation to global warming. Such environmental changes would add to the impact of fish extraction by the industrial fisheries on the southern food webs.     

Influence of local and external processes on the annual nitrogen cycle and primary productivity on Georges Bank: A 3-D biological–physical modeling study

Georges Bank is one of the world's most highly productive marine areas, but the mechanisms of nutrient supply to support such high productivity remain poorly understood. Intrusions of nutrient-poor Labrador Slope Water (LSW) into the Gulf of Maine (NAO-dependent) potentially can reduce nutrient delivery to the bank, but this mechanism has not been quantitatively examined. In this paper, we present the first whole-year continuous model simulation results using a biological–physical model developed for the Gulf of Maine/Georges Bank region. This high-resolution three-dimensional coupled model consists of the Finite Volume Coastal Ocean Model (FVCOM) and a Nitrogen–Phytoplankton–Zooplankton–Detritus (NPZD) model, and was used to examine the influences of local and external processes on nitrogen and phytoplankton dynamics on Georges Bank. The model captured the general pattern of spatial-temporal distributions of nitrogen and phytoplankton and provided a diagnostic analysis of different processes that control nitrogen fluxes on Georges Bank. Specifically, numerical experiments were conducted to examine seasonal variation in nitrogen transport into the central bank (new nitrogen supply) versus nitrogen regenerated internally in this region. Compared with previous observation-based studies, the model provided a quantitative estimate of nitrogen flux by integrating the transport over a longer time period and a complete spatial domain. The results suggest that, during summer months, internal nitrogen regeneration is the major nitrogen source for primary production on the central bank, while nitrogen supply through physical transport (e.g. tidal pumping) contributes about 1/5 of the total nitrogen demand, with an estimated on-bank nitrogen transport at least 50% less than previous estimates. By comparing the model runs using different nitrogen concentrations in deep Slope Water, the potential influence of NAO-dependent intrusions of LSW was examined. The results suggest that the change of nitrogen concentration in the deep Slope Water may not have a significant impact on nitrogen and phytoplankton dynamics on the well-mixed central bank, largely due to limited nutrient exchange across the tidal mixing front and enhanced near-frontal nutrient uptake. However, relatively more significant impact was observed in the model simulations if both well-mixed and seasonally-stratified areas (inside 100 m isobath of the bank) were considered in flux calculations.     

A generalized model of pelagic biogeochemistry for the global ocean ecosystem. Part II: Numerical simulations

This paper presents a global ocean implementation of a multi-component model of marine pelagic biogeochemistry coupled on-line with an ocean general circulation model forced with climatological surface fields (PELAgic biogeochemistry for Global Ocean Simulations, PELAGOS). The final objective is the inclusion of this model as a component in an Earth System model for climate studies. The pelagic model is based on a functional stoichiometric representation of marine biogeochemical cycles and allows simulating the dynamics of C, N, P, Si, O and Fe taking into account the variation of their elemental ratios in the functional groups. The model also includes a parameterization of variable chlorophyll/carbon ratio in phytoplankton, carrying chl as a prognostic variable. The first part of the paper analyzes the contribution of non-local advective–diffusive terms and local vertical processes to the simulated chl distributions. The comparison of the three experiments shows that the mean chl distribution at higher latitudes is largely determined by mixing processes, while vertical advection controls the distribution in the equatorial upwelling regions. Horizontal advective and diffusive processes are necessary mechanisms for the shape of chl distribution in the sub-tropical Pacific. In the second part, the results have been compared with existing datasets of satellite-derived chlorophyll, surface nutrients, estimates of phytoplankton community composition and primary production data. The agreement is reasonable both in terms of the spatial distribution of annual means and of the seasonal variability in different dynamical oceanographic regions. Results indicate that some of the model biases in chl and surface nutrients distributions can be related to deficiencies in the simulation of physical processes such as advection and mixing. Other discrepancies are attributed to inadequate parameterizations of phytoplankton functional groups. The model has skill in reproducing the overall distribution of large and small phytoplankton but tends to underestimate diatoms in the northern higher latitudes and overestimate nanophytoplankton with respect to picoautotrophs in oligotrophic regions. The performance of the model is discussed in the context of its use in climate studies and an approach for improving the parameterization of functional groups in deterministic models is outlined.     

An integrated water/sediment approach to study plankton (a case study in the southern Adriatic Sea)

In marine coastal areas many planktonic species produce resting stages (cysts) that sink to the bottom. Integrated sampling from both the water column (to collect active stages), and sediments (to collect cysts), could be useful to achieve more complete information about plankton composition.In the framework of the “INTERREG II Albania-Italy Project” an oceanographic survey was carried out aboard the r/v “Italica” from 20 to 31 October 2000. The survey interested the northern Albanian coast (Gulf of Drin) and the northern Apulian coast (Gulf of Manfredonia) on the opposite sides of the South Adriatic Sea. The plankton was collected from 14 stations. A total of 188 categories were recognized in plankton samples. Among those categories, 130 species were recognized (87 of phytoplankton, 43 of microzooplankton), and only 53 (40.8%) resulted common to both the Adriatic sides. A total of 69 cyst morphotypes were recognized in sediment samples; 38 of them were classified at the level of species. A statistical analysis of the micro-zooplankton species abundance showed a segregation of the two areas better than that obtained with the phytoplankton. Cyst distribution in the sediments showed a good gulf-segregation too. In addition, they allowed us to find complementary information, particularly for dinoflagellates. The most abundant species in the water column were not equally dominant as resting stages in the sediments. Sediment sampling allowed further information about the composition of the plankton communities, and suggested us to search for a new method to enhance the yield of less abundant cysts.     

Large-scale physical controls on phytoplankton growth in the Irminger Sea, Part II: Model study of the physical and meteorological preconditioning

The upper water column in the Irminger Sea is characterized by cold fresh arctic and subarctic waters and warm saline North Atlantic waters. In this study the local physical and meteorological preconditioning of the phytoplankton development over an annual cycle in the upper water column in four physical zones of the Irminger Sea is investigated. Data from four cruises of the UK's Marine Productivity programme are combined with results from a coupled biological–physical nitrogen–phytoplankton–zooplankton–detritus model run using realistic forcing. The observations and model predictions are compared and analyzed to identify the key parameters and processes which determine the observed heterogeneity in biological production in the Irminger Sea. The simulations show differences in the onset of the bloom, in the time of the occurrence of the maximum phytoplankton biomass and in the length of the bloom between the zones. The longest phytoplankton bloom of 90 days duration was predicted for the East Greenland Current of Atlantic origin zone. In contrast, for the Central Irminger Sea zone a phytoplankton bloom with a start at the beginning of May and the shortest duration of only 70 days was simulated. The latest onset of the phytoplankton bloom in mid May and the latest occurrence of the maximum biomass (end of July) were predicted for the Northern Irminger Current zone. Here the bloom lasted for 80 days. In contrast the phytoplankton bloom in the Southern Irminger Current zone started at the same time as in Central Irminger Sea, but peaked end of June and lasted for 80 days. For all four zones relatively low daily (0.3–0.5 g C m^− 2d^− 1) and annual primary production was simulated, ranging between 35.6 g C m^− 2y^− 1 in the East Greenland Current of Atlantic origin zone and 45.6 g C m^− 2y^− 1 in the Northern Irminger Current zone. The model successfully simulated the observed regional and spatial differences in terms of the maximum depth of winter mixing, the onset of stratification and the development of the seasonal thermocline, and the differences in biological characteristics between the zones. The initial properties of the water column and the seasonal cycle of physical and meteorological forcing in each of the zones are responsible for the observed differences during the Marine Productivity cruises. The timing of the transition from mixing to stratification regime, and the different prevailing light levels in each zone are identified as the crucial processes/parameters for the understanding of the dynamics of the pelagic ecosystem in the Irminger Sea.     

Effects of small-scale turbulence on the growth of two diatoms of different size in a phosphorus-limited medium

The effect of turbulence on the nutrient flux towards osmotrophic cells is predicted to be size dependent. This should translate into growth. We experimentally followed and modelled the growth of two marine diatoms of different size (Thalassiosira pseudonana, 6 μm in diameter and Coscinodiscus sp., ca. 109 μm in diameter) under still water and turbulent conditions, using a shaker table. Experiments were done with phosphorus-limited cultures and lasted for ca. 5 days. Turbulence enhanced the growth of Coscinodiscus sp. in agreement with theory but not the growth of T. pseudonana, which was actually slightly lower under turbulence. At the end of the experiments there were about 1.7 times as many Coscinodiscus sp. cells in the turbulent treatment than in the still treatment, while for T. pseudonana almost the same cell concentration was found in both conditions. In addition, the Coscinodiscus sp. cells growing under still conditions presented a higher specific alkaline phosphatase activity than those growing in turbulence which indicates a higher need for phosphorus in the still cultures. A simple dynamic model, based on Michaelis–Menten nutrient uptake kinetics, needed nearly no optimisation other than using observed initial conditions of phosphate and cell concentrations. The model showed how an increased nutrient flux towards the cells translates non-linearly into cell growth, most likely by affecting the half-saturation constant (K_M). However, since Coscinodiscus sp. experienced significant mortality and cells partially settled to the bottom of the containers, unequivocal support for the size-dependent effect of turbulence on nutrient uptake will require further experiments and more sophisticated modelling. The mechanisms to connect an increased nutrient flux towards cells with population growth and whether this process is size dependent are important in parameterizing the effects of turbulence on marine plankton in coupled physical–biological models.     

Study of the seasonal cycle of the biogeochemical processes in the Ligurian Sea using a 1D interdisciplinary model

A one-dimensional coupled physical–biogeochemical model has been built to study the pelagic food web of the Ligurian Sea (NW Mediterranean Sea). The physical model is the turbulent closure model (version 1D) developed at the GeoHydrodynamics and Environmental Laboratory (GHER) of the University of Liège. The ecosystem model contains 19 state variables describing the carbon and nitrogen cycles of the pelagic food web. Phytoplankton and zooplankton are both divided in three size-based compartments and the model includes an explicit representation of the microbial loop including bacteria, dissolved organic matter, nano-, and microzooplankton. The internal carbon/nitrogen ratio is assumed variable for phytoplankton and detritus, and constant for zooplankton and bacteria. Silicate is considered as a potential limiting nutrient of phytoplankton's growth. The aggregation model described by Kriest and Evans in (Proc. Ind. Acad. Sci., Earth Planet. Sci. 109 (4) (2000) 453) is used to evaluate the sinking rate of particulate detritus. The model is forced at the air–sea interface by meteorological data coming from the “Côte d'Azur” Meteorological Buoy. The dynamics of atmospheric fluxes in the Mediterranean Sea (DYFAMED) time-series data obtained during the year 2000 are used to calibrate and validate the biological model. The comparison of model results within in situ DYFAMED data shows that although some processes are not represented by the model, such as horizontal and vertical advections, model results are overall in agreement with observations and differences observed can be explained with environmental conditions.     

Intrusions of eastern North Atlantic central waters and phytoplankton in the north and northwestern Iberian shelf during spring

The influence of intrusions of eastern North Atlantic central water (ENACW) in the north and northwestern Iberian shelf on phytoplankton composition and abundance and on particle-size distributions of seston was analyzed using data collected on three extensive cruises during spring 1991 and 1992. Water with temperature and salinity values between 12.20 and 13.86 °C and between 35.66 and 35.98 psu, respectively, characteristics of the subtropical type of ENACW (ENACW_t), was detected in the upper 100 m of the water-column in all cruises, but particularly in the western coast in 1992. The highest salinity values of this water were found near the surface (0–100-m depth) and in early spring 1992, while minimum salinity values, and also minimum geographical extension, were found in late spring in both years. Phytoplankton blooms concentrated in frontal areas between different water types, with maximum intensity and extension in early spring.Using temperature and salinity characteristics, samples were classified in four groups corresponding to the major water types found in the region: Bay of Biscay central water (BBCW), two segments of ENACW of different salinity and surface water influenced by continental runoff. This classification was significantly confirmed by three independent discriminant analyses using hydrographic and chemical (dissolved nutrients and chlorophyll) variables, phytoplankton species abundance variables and particle-size concentration of seston variables. Phytoplankton blooms related to the presence of saline waters were characterized by the dominance of either chain-forming diatoms or a mixture of diatoms and phytoflagellates and high concentrations of seston. The diatom species dominating in saline waters were typical of upwelling-induced blooms occurring generally during summer. Blooms occurring in waters influenced by runoff also contained diatoms but in lower numbers than those of saline waters. Nutrients were not exhausted in the region, suggesting that phytoplankton populations were still in active growth. These results are interpreted taking into account the known variability in water-mass formation and in the poleward current driving ENACW_t along the shelf, and indicate that saline intrusions are a major feature affecting the distribution and composition of plankton in the spring in the southern Bay of Biscay, thus enlarging to a wider spatial scale their reported influence on the pelagic ecosystem.     

Wind-triggered events of phytoplankton downward flux in the Northeast Water Polynya

Phytoplankton carbon fluxes were studied in the Northeast Water (NEW) Polynya, off the eastern coast of Greenland (79° to 81°N, 6° to 17°W), during summer 1993. The downward flux of organic particles was determined during 54 days using a sediment trap moored at a fixed location, below the pycnocline (130 m). The hypothesis of the present study is that wind events were ultimately responsible for the events of diatoms downward flux recorded in the trap.Wind conditions can influence the vertical transport of phytoplankton by affecting (1) the environmental conditions (e.g. hydrostatic pressure, nutrient concentrations, and irradiance) encountered by phytoplankton during their vertical excursion, and (2) the aggregation and disaggregation of phytoplankton flocs. The first mechanism affects the physiological regulation of buoyancy, whereas the second one affects the size and shape of settling particles.Using field data (wind velocity, density profiles and phytoplankton abundance), we assessed the potential aggregation and the vertical excursion of phytoplankton in surface waters. The results show that, upstream from the trap, wind and hydrodynamic conditions were sometimes favourable to the downward export of phytoplankton. Lag-correlation between time series of wind and phytoplankton downward flux shows that flux events lagged wind events by ca. 16 days. Given that the average current velocity in the top 100 m was ca. 10 cm s⁻¹, a lag of 16 days corresponded to a lateral transport of ca. 130 km, upstream from the sediment trap, where phytoplankton production was lower than at the location of the trap. According to that scenario, 21% to 60% of primary production was exported to depth during wind events. If we had assumed instead a tight spatial coupling between the material collected in the trap and the relatively high phytoplankton production at the location of the trap, we would have concluded that <7% of primary production was exported to depth. The difference between the two scenarios has great implications for the fate of phytoplankton. Our results stress the importance of investigating the spatial coupling between surface and trap data before assessing the pathways of phytoplankton carbon cycling.     

Lagrangian modelling of plankton motion: From deceptively simple random walks to Fokker–Planck and back again

The movement of plankton, either by turbulent mixing or their own inherent motility, can be simulated in a Lagrangian framework as a random walk. Validation of random walk simulations is essential. There is a continuum of mathematically valid stochastic integration schemes upon which random walk simulations depend, each of which lead to radically different macro-scale dynamics as expressed in their corresponding Fokker–Planck equations. In addition, diffusivity is not a unique parameter describing a random walk and its corresponding Fokker–Planck equation. Spatially varying translation speed and turn frequency have different effects on population distributions. Validation requires extra information either in the form of the well-mixed condition for physical diffusion, or in detailed information on the sensing ability, internal state modulation and swimming response for plankton motility.