Optimizing Models of the North Atlantic Spring Bloom Using Physical, Chemical and Bio-Optical Observations from a Lagrangian Float

The North Atlantic spring bloom is one of the main events that lead to carbon export to the deep ocean and drive oceanic uptake of CO(2) from the atmosphere. Here we use a suite of physical, bio-optical and chemical measurements made during the 2008 spring bloom to optimize and compare three different models of biological carbon export. The observations are from a Lagrangian float that operated south of Iceland from early April to late June, and were calibrated with ship-based measurements. The simplest model is representative of typical NPZD models used for the North Atlantic, while the most complex model explicitly includes diatoms and the formation of fast sinking diatom aggregates and cysts under silicate limitation. We carried out a variational optimization and error analysis for the biological parameters of all three models, and compared their ability to replicate the observations. The observations were sufficient to constrain most phytoplankton-related model parameters to accuracies of better than 15 %. However, the lack of zooplankton observations leads to large uncertainties in model parameters for grazing. The simulated vertical carbon flux at 100 m depth is similar between models and agrees well with available observations, but at 600 m the simulated flux is larger by a factor of 2.5 to 4.5 for the model with diatom aggregation. While none of the models can be formally rejected based on their misfit with the available observations, the model that includes export by diatom aggregation has a statistically significant better fit to the observations and more accurately represents the mechanisms and timing of carbon export based on observations not included in the optimization. Thus models that accurately simulate the upper 100 m do not necessarily accurately simulate export to deeper depths.

Plankton biomass, POC and PON concentrations, and sinking velocities during the 2008 spring diatom bloom in Disko Bay

Phytoplankton and copepod succession was investigated in Disko Bay, western Greenland from February to July 2008. The spring phytoplankton bloom developed immediately after the breakup of sea ice and reached a peak concentration of 24 mg chl a/m**3 2 wk later. The bloom was analyzed during 3 phases: the developing, the decaying, and the post-bloom phases. Grazing impact by the copepod community was assessed by 4 methods; gut fluorescence, in situ faecal pellet production, and egg and faecal pellet production from bottle incubations. Calanus spp. dominated the mesozooplankton community. They were present from the initiation of the bloom but only had a small grazing impact on the phytoplankton. Consequently, there was a close coupling between the spring phytoplankton bloom and sedimentation of particulate organic carbon (POC). Out of 1836 ±180 mg C/m**2/d leaving the upper 50 m, 60 % was phytoplankton based carbon (PPC). The composition and quality of the sedimenting material changed throughout the bloom succession from PPC dominance in the initial phase with a POC/PON ratio close to 6.6 to a dominance of amorphous detritus with a higher POC/PON ratio (>10) in the post-bloom phase. The succession and fate of the phytoplankton spring bloom was controlled by nitrogen limitation and subsequent sedimentation, while grazing-mediated flux by the Calanus-dominated copepod community played a minor role in the termination of the spring bloom of Disko Bay.

Interannual Variability in Timing of Bloom Initiation in the California Current System

In the California Current System the spring transition from poleward to equatorward alongshore wind stress heralds the beginning of upwelling-favorable conditions. The phytoplankton response to this transition is investigated using 8 years ( 1998-2005) of daily, 4-km resolution, Sea-viewing Wide Field of view Sensor ( SeaWiFS) chlorophyll a concentration data. Cluster analysis of the chlorophyll a time series at each location is used to separate the inshore upwelling region from offshore and oligotrophic areas. An objective method for estimating the timing of bloom initiation is used to construct a map of the mean bloom start date. Interannual variability in bloom timing and magnitude is investigated in four regions: 45 degrees N - 50 degrees N, 40 degrees N - 45 degrees N, 35 degrees N - 40 degrees N and 20 degrees N - 35 degrees N. Daily satellite derived wind data ( QuikSCAT) allow the timing of the first episode of persistently upwelling favorable winds to be estimated. Bloom initiation generally coincides with the onset of upwelling winds ( +/- 15 days). South of similar to 35 degrees N, where winds are southward year-round, the timing of increased chlorophyll concentration corresponds closely to timing of the seasonal increase in upwelling intensity. A 1-D model and satellite derived photosynthetically available radiation data are used to estimate time series of depth- averaged irradiance. In the far north of the region (> 46 degrees N) light is shown to limit phytoplankton growth in early spring. In 2005 the spring bloom in the northern regions (> 35 degrees N) had a "false start''. A sharp increase in chl a in February quickly receded, and a sustained increase in biomass was delayed until July. We hypothesize that this resulted in a mismatch in timing of food availability to higher trophic levels.

(Table 2) Concentration of phytoplankton pigments in Adventfjorden in spring and summer of 1997

The study was carried out from April 30 until July 13 of 1997 in Adventfjorden (Spitsbergen). Formation of a less saline and warmer surface water (~1 m thick) caused by melting of the ice was observed in the fjord during the first days of May. In summer the less saline surface layer was about 3 m thick. Euphotic depth measured under the ice sheet reached 12 m, whereas load of mineral matter brought with riverine discharge in summer (content of total particulate matter in the fjord reached 1.66 kg/m**2) dramatically reduced euphotic zone depth to 0.35 m. By pigment measurement three phases of phytoplankton development in Adventfjorden were distinguished: (1) spring bloom that has started under fast ice and reached maximum in the mid of May, (2) stagnation period in June, (3) increase of pigment concentration in July, what could indicate start of the next algae bloom. Analyses of chlorophylls and carotenoids revealed that diatoms (chl c, fucoxanthin), and green algae (chl b, lutein) dominated phytoplankton community in the fjord. Moreover, presence of peridinin indicates presence of Dinophyta and alloxanthin - occurence of Cryptophyta. In May and June 1997 phytoplankton appeared mainly in the surface of water, while in July, as a result of inflow of turbulent riverine waters into Adventfjorden, algae cells were pushed down and the highest numbers were observed at depth ~20 m. Great phaeopigments to chl a ratio (= 0.54) found in fjord seston in June and July probably shows strong impact of zooplankton grazing on phytoplankton development. High contribution of chlorophyllide a in porphyrin a poll in samples collected under fast ice (chlorophyllide a / chl a ratio = 0.18) reflects the final stage of algal communitie succession in ice, just before spring ice melt and release of biota to oceanic water. Chlorophyllide a content during summer was minor or not detectable, demonstrating that diatom cells were in good physiological condition. High chl a allomer / chl a ratio (average = 0.11 for the period investigated) confirms high oxygen concentration in environment of Adventfjorden.

Climatic patterns and physical controls of chlorophyll-a in the Northeast Atlantic

Tese de Doutoramento, Física, 17 de Dezembro de 2013, Universidade dos Açores.

External forcings, oceanographic processes and particle flux dynamics in Cap de Creus submarine canyon, NW Mediterranean Sea

Particle fluxes (including major components and grain size), and oceanographic parameters (near-bottom water temperature, current speed and suspended sediment concentration) were measured along the Cap de Creus submarine canyon in the Gulf of Lions (GoL; NW Mediterranean Sea) during two consecutive winter-spring periods (2009 2010 and 2010 2011). The comparison of data obtained with the measurements of meteorological and hydrological parameters (wind speed, turbulent heat flux, river discharge) have shown the important role of atmospheric forcings in transporting particulate matter through the submarine canyon and towards the deep sea. Indeed, atmospheric forcing during 2009 2010 and 2010 2011 winter months showed differences in both intensity and persistence that led to distinct oceanographic responses. Persistent dry northern winds caused strong heat losses (14.2 × 103 W m−2) in winter 2009 2010 that triggered a pronounced sea surface cooling compared to winter 2010 2011 (1.6 × 103 W m−2 lower). As a consequence, a large volume of dense shelf water formed in winter 2009 2010, which cascaded at high speed (up to ∼1 m s−1) down Cap de Creus Canyon as measured by a current-meter in the head of the canyon. The lower heat losses recorded in winter 2010 2011, together with an increased river discharge, resulted in lowered density waters over the shelf, thus preventing the formation and downslope transport of dense shelf water. High total mass fluxes (up to 84.9 g m−2 d−1) recorded in winter-spring 2009 2010 indicate that dense shelf water cascading resuspended and transported sediments at least down to the middle canyon. Sediment fluxes were lower (28.9 g m−2 d−1) under the quieter conditions of winter 2010 2011. The dominance of the lithogenic fraction in mass fluxes during the two winter-spring periods points to a resuspension origin for most of the particles transported down canyon. The variability in organic matter and opal contents relates to seasonally controlled inputs associated with the plankton spring bloom during March and April of both years.

Climate driven changes in temperature, pH and food quality : effects on copepod reproduction

Increased emissions of greenhouse gases into the atmosphere are causing an anthropogenic climate change. The resulting global warming challenges the ability of organisms to adapt to the new temperature conditions. However, warming is not the only major threat. In marine environments, dissolution of carbon dioxide from the atmosphere causes a decrease in surface water pH, the so called ocean acidification. The temperature and acidification effects can interact, and create even larger problems for the marine flora and fauna than either of the effects would cause alone. I have used Baltic calanoid copepods (crustacean zooplankton) as my research object and studied their growth and stress responses using climate predictions projected for the next century. I have studied both direct temperature and pH effects on copepods, and indirect effects via their food: the changing phytoplankton spring bloom composition and toxic cyanobacterium. The main aims of my thesis were: 1) to find out how warming and acidification combined with a toxic cyanobacterium affect copepod reproductive success (egg production, egg viability, egg hatching success, offspring development) and oxidative balance (antioxidant capacity, oxidative damage), and 2) to reveal the possible food quality effects of spring phytoplankton bloom composition dominated by diatoms or dinoflagellates on reproducing copepods (egg production, egg hatching, RNA:DNA ratio). The two copepod genera used, Acartia sp. and Eurytemora affinis are the dominating mesozooplankton taxa (0.2 – 2 mm) in my study area the Gulf of Finland. The 20°C temperature seems to be within the tolerance limits of Acartia spp., because copepods can adapt to the temperature phenotypically by adjusting their body size. Copepods are also able to tolerate a pH decrease of 0.4 from present values, but the combination of warm water and decreased pH causes problems for them. In my studies, the copepod oxidative balance was negatively influenced by the interaction of these two environmental factors, and egg and nauplii production were lower at 20°C and lower pH, than at 20°C and ambient pH. However, presence of toxic cyanobacterium Nodularia spumigena improved the copepod oxidative balance and helped to resist the environmental stress, in question. In addition, adaptive maternal effects seem to be an important adaptation mechanism in a changing environment, but it depends on the condition of the female copepod and her diet how much she can invest in her offspring. I did not find systematic food quality difference between diatoms and dinoflagellates. There are both good and bad diatom and dinoflagellate species. Instead, the dominating species in the phytoplankton bloom composition has a central role in determining the food quality, although copepods aim at obtaining as a balanced diet as possible by foraging on several species. If the dominating species is of poor quality it can cause stress when ingested, or lead to non-optimal foraging if rejected. My thesis demonstrates that climate change induced water temperature and pH changes can cause problems to Baltic Sea copepod communities. However, their resilience depends substantially on their diet, and therefore the response of phytoplankton to the environmental changes. As copepods are an important link in pelagic food webs, their future success can have far reaching consequences, for example on fish stocks.

Phytoplankton Dynamics in Three Rocky Mountain Lakes, Colorado, U.S.A.

In 1984 and 1985 seasonal changes in phytoplankton were studied in a system of three lakes in Loch Vale, Rocky Mountain National Park, Colorado. Three periods were evident: (1) A spring bloom, during snowmelt, of the planktonic diatom Asterionella Formosa, (2) a mid- summer period of minimal algal abundance, and (3) a fall bloom of the blue-green alga Oscillatoria limnetica. Seasonal phytoplankton dynamics in these lakes are controlled partially by the rapid flushing rate during snowmelt and the transport of phytoplankton from the highest lake to the lower lakes by the stream, Icy Brook. During snowmelt, the A. formosa population in the most downstream lake has a net rate of increase of 0.34 d-1, which is calculated from the flushing rate and from the A. formosa abundance in the inflow from the upstream lake and in the downstream lake. Measurement of photosynthetic rates at different depths during the three periods confirmed the rapid growth of A. formosa during the spring. The decline in A. formosa after snowmelt may be related to grazing by developing zooplankton populations. The possible importance of the seasonal variations in nitrate concentrations were evaluated in situ enrichment experiments. For A. formosa and O. limnetica populations, growth stimulation resulted from 8- or 16-micromolar amendments of calcium nitrate and sulfuric acid, but the reason for this stimulation could not be determined from these experiments.

Bio-physical interactions and feedbacks in a global climate model

This PhD thesis addresses the topic of large-scale interactions between climate and marine biogeochemistry. To this end, centennial simulations are performed under present and projected future climate conditions with a coupled ocean-atmosphere model containing a complex marine biogeochemistry model. The role of marine biogeochemistry in the climate system is first investigated. Phytoplankton solar radiation absorption in the upper ocean enhances sea surface temperatures and upper ocean stratification. The associated increase in ocean latent heat losses raises atmospheric temperatures and water vapor. Atmospheric circulation is modified at tropical and extratropical latitudes with impacts on precipitation, incoming solar radiation, and ocean circulation which cause upper-ocean heat content to decrease at tropical latitudes and to increase at middle latitudes. Marine biogeochemistry is tightly related to physical climate variability, which may vary in response to internal natural dynamics or to external forcing such as anthropogenic carbon emissions. Wind changes associated with the North Atlantic Oscillation (NAO), the dominant mode of climate variability in the North Atlantic, affect ocean properties by means of momentum, heat, and freshwater fluxes. Changes in upper ocean temperature and mixing impact the spatial structure and seasonality of North Atlantic phytoplankton through light and nutrient limitations. These changes affect the capability of the North Atlantic Ocean of absorbing atmospheric CO2 and of fixing it inside sinking particulate organic matter. Low-frequency NAO phases determine a delayed response of ocean circulation, temperature and salinity, which in turn affects stratification and marine biogeochemistry. In 20th and 21st century simulations natural wind fluctuations in the North Pacific, related to the two dominant modes of atmospheric variability, affect the spatial structure and the magnitude of the phytoplankton spring bloom through changes in upper-ocean temperature and mixing. The impacts of human-induced emissions in the 21st century are generally larger than natural climate fluctuations, with the phytoplankton spring bloom starting one month earlier than in the 20th century and with ~50% lower magnitude. This PhD thesis advances the knowledge of bio-physical interactions within the global climate, highlighting the intrinsic coupling between physical climate and biosphere, and providing a framework on which future studies of Earth System change can be built on.

Springtime Nutrient and Phytoplankton Dynamics on Georges Bank

The dynamics of phytoplankton and nutrients before, during and after the winter-spring bloom on Georges Bank were studied on 6 monthly survey cruises from January to June 1999. We measured hydrography, phytoplankton cell densities, chlorophyll a, dissolved inorganic nutrients (NO3 + NO2, NH4, Si(OH)(4), PO4), dissolved organic nitrogen (DON) and phosphorus (DOP), particulate organic carbon (POC) and nitrogen (PON) and total particulate phosphorus (TPP). We present evidence that phytoplankton production may be significant year-round, and that the winter-spring bloom may have started in January. From January to April the phytoplankton was comprised almost exclusively of diatoms, reaching cell densities in March and April of ca. 450 cells ml(-1); chlorophyll a concentrations exceeded 10 mug l(-1) in April. Diatoms decreased to relatively low levels in May (< 50 x 10(3) cells l(-1)) and increased again in June (>300 x 10(3) cells l(-1)). Densities of dinoflagellates and nanoflagellates were low (< 10 x 10(3) cells l(-1)) from January to April, and increased in May and June to nearly 300 x 10(3) cells l(-1). Nitrate + nitrite concentrations in January were <3 muM in the shallow, central portion of the bank and decreased steadily each month. Silicate was also <3 muM over an even larger area of the central bank in January and declined to <1.5 muM over most of the Bank in April. The data suggest that silicate depletion, not DIN, contributed to the cessation of the diatom bloom. Regeneration of silicate occurred in May and June, presumably as a result of rising water temperatures in late spring which increased the dissolution rate of diatom frustules from the earlier diatom bloom. Dissolved organic nitrogen may have been utilized at the start of the winter-spring bloom; concentrations were ca, 14 muM in January, dropping to < 6 mug l(-1) in February, after which DON concentrations steadily rose to > 15 mug l(-1) in June. Overall micro-and nanoplankton biomass, measured as POC, PON and TPP, increased over the 6 mo period, as did nutritional quality of that biomass as indicated by declining C:N ratios. Our results suggest there may have been an increase in the heterotrophic component of the plankton in May and June which coincided with a second burst in diatom abundance. We discuss general features of planktonic production and nutrient dynamics with respect to year-round production on the Bank.

An individual based model spatial distribution, production and biomass for Calanus finmarchicus in the Norwegian Sea

The copepod Calanus finmarchicus is the dominant species of the meso-zooplankton in the Norwegian Sea, and constitutes an important link between the phytoplankton and the higher trophic levels in the Norwegian Sea food chain. An individualbased model for C. finmarchicus, based on super-individuals and evolving traits for behaviour, stages, etc., is two-way coupled to the NORWegian ECOlogical Model system (NORWECOM). One year of modelled C. finmarchicus spatial distribution, production and biomass are found to represent observations reasonably well. High C. finmarchicus abundance is found along the Norwegian shelf-break in the early summer, while the overwintering population is found along the slope and in the deeper Norwegian Sea basins. The timing of the spring bloom is generally later than in the observations. Annual Norwegian Sea production is found to be 29 million tonnes of carbon and a production to biomass (P/B) ratio of 4.3 emerges. Sensitivity tests show that the modelling system is robust to initial values of behavioural traits and with regards to the number of super-individuals simulated given that this is above about 50,000 individuals. Experiments with the model system indicate that it provides a valuable tool for studies of ecosystem responses to causative forces such as prey density or overwintering population size. For example, introducing C. finmarchicus food limitations reduces the stock dramatically, but on the other hand, a reduced stock may rebuild in one year under normal conditions. The NetCDF file contains model grid coordinates and bottom topography.