Simulation of upper-ocean biogeochemistry with a flexible-composition phytoplankton model: C, N and Si cycling in the western Sargasso Sea

We report the first application of a biogeochemical model in which the major elemental composition of the phytoplankton is flexible, and responds to changing light and nutrient conditions. The model includes two phytoplankton groups: diatoms and non-siliceous picoplankton. Both fix C in accordance with photosynthesis-irradiance relationships used in other models and take up NO₃⁻ and NH₄⁺ (and Si(OH)₄ for diatoms) following Michaelis-Menten kinetics. The model allows for light dependence of photosynthesis and NO₃⁻ uptake, and for the observed near-total light independence of NH₄⁺ uptake and Si(OH)₄ uptake. It tracks the resulting C/N ratios of both phytoplankton groups and Si/N ratio of diatoms, and permits uptake of C, N and Si to proceed independently of one another when those ratios are close to those of nutrient-replete phytoplankton. When the C/N or Si/N ratio of either phytoplankton group indicates that its growth is limited by N, Si or light, uptake of non-limiting elements is controlled by the content of the limiting element in accordance with the cell-quota formulation of Droop (J. Mar. Biol. Ass. U.K 54 (1974) 825).We applied this model to the Bermuda Atlantic Time-series Study (BATS) site in the western Sargasso Sea. The model was tuned to produce vertical profiles and time courses of [NO₃⁻], [NH₄⁺] and [Si(OH)₄] that are consistent with the data, by adjusting the kinetic parameters for N and Si uptake and the rate of nitrification. The model then reproduces the observed time courses of chlorophyll-a, particulate organic carbon and nitrogen, biogenic silica, primary productivity, biogenic silica production and POC export with no further tuning. Simulated C/N and Si/N ratios of the phytoplankton indicate that N is the main growth-limiting nutrient throughout the thermally stratified period and that [Si(OH)₄], although always limiting to the rate of Si uptake by diatoms, seldom limits their growth rate. The model requires significant nitrification in the upper 200 m to yield realistic time courses and vertical profiles of [NH₄⁺] and [NO₃⁻], suggesting that NO₃⁻ is not supplied to the upper water column entirely by physical processes. A nitrification-corrected f-ratio (f_NC), calculated for the upper 200 m as: (NO₃⁻ uptake—nitrification)/(NO₃⁻ uptake+NH₄⁺ uptake) has annual values ranging from only 0.05–0.09, implying that 90–95% of the N taken up annually by phytoplankton is supplied by biological regeneration (including nitrification) in the upper 200 m. Reported discrepancies between estimates of organic C export based on seasonal chemical changes and POC export measured at the BATS site can be almost completely resolved if there is significant regeneration of NO₃⁻ via organic-matter decomposition in the upper 200 m.     

Summer phytoplankton blooms in the oligotrophic North Pacific Subtropical Gyre: Historical perspective and recent observations

The export of organic matter from the oceanic euphotic zone is a critical process in the global biogeochemical cycling of bioelements (C, N, P, Si). Much of this export occurs in the form of sinking particles, which rain down into the unlit waters of the deep sea. Classical models of oceanic production and export balance this gravitational loss of particulate bioelements with an upward flux of dissolved nutrients, and they describe reasonably well those areas of the ocean where deep winter mixing occurs. The surface waters of the North Pacific Subtropical Gyre (NPSG), however, are strongly stratified and chronically nutrient-depleted, especially in summer. Nevertheless, there is ample evidence that blooms of phytoplankton and subsequent pulses of particle export occur during the height of summer stratification in these waters, especially to the northeast of the Hawaiian Islands. These blooms impact regional bioelemental cycling and act as a food source to the deep-sea benthos. We review here numerous published observations of these events in the NPSG, and present new data collected at Station ALOHA (22.75°N, 158°W) during the first 176 cruises of the Hawaii Ocean Time-series program (1988-2005), along with results from transect cruises conducted in the region in 1996 and 2005. We suggest that the summer phytoplankton bloom can be considered a frequent, perhaps annual feature in the northeastern NPSG, and that its perceived stochastic nature is a manifestation of chronic undersampling in time and space. The bloom is typically dominated by only a few genera of large diatoms and the cyanobacterium Trichodesmium. It appears to be consistently supported by dinitrogen fixation, but the fate of the organic matter produced during the summer depends critically on the species composition of the responsible diazotrophs. We estimate that the summer bloom is responsible for up to 38% of N₂ fixation and up to 18% of N-based new production annually at Station ALOHA. We hypothesize that the spatial distribution, timing and magnitude of the bloom may be determined largely by the physical and biological processes controlling new phosphorus delivery into the euphotic zone during the summer and the preceding winter.     

Modelling phytoplankton succession on the Bering Sea shelf: role of climate influences and trophic interactions in generating Emiliania huxleyi blooms 1997–2000

Several years of continuous physical and biological anomalies have been affecting the Bering Sea shelf ecosystem starting from 1997. Such anomalies reached their peak in a striking visual phenomenon: the first appearance in the area of bright waters caused by massive blooms of the coccolithophore Emiliania huxleyi (E. huxleyi). This study is intended to provide an insight into the mechanisms of phytoplankton succession in the south-eastern part of the shelf during such years and addresses the causes of E. huxleyi success by means of a 2-layer ecosystem model, field data and satellite-derived information. A number of potential hypotheses are delineated based on observations conducted in the area and on previous knowledge of E. huxleyi general ecology. Some of these hypotheses are then considered as causative factors and explored with the model. The unusual climatic conditions of 1997 resulted most notably in a particularly shallow mixed layer depth and high sea surface temperature (about 4 °C above climatological mean). Despite the fact that the model could not reproduce for E. huxleyi a clear non-bloom to bloom transition (pre- vs. post-1997), several tests suggest that this species was favoured by the shallow mixed layer depth in conjunction with a lack of photoinhibition. A top-down control by microzooplankton selectively grazing phytoplankton other than E. huxleyi appears to be responsible for the long persistence of the blooms. Interestingly, observations reveal that the high N:P ratio hypothesis, regarded as crucial in the formation of blooms of this species in previous studies, does not hold on the Bering Sea shelf.     

Modelling the influence of environmental factors on the physiological status of the Pacific oyster Crassostrea gigas in an estuarine embayment; The Baie des Veys (France)

It is well known that temporal changes in bivalve body mass are strongly correlated with temporal variations in water temperature and food supply. In order to study the influence of the year-to-year variability of environmental factors on oyster growth, we coupled a biogeochemical sub-model, which simulates trophic resources of oysters (i.e. phytoplankton biomass via chlorophyll a), and an ecophysiological sub-model, which simulates growth and reproduction (i.e. gametogenesis and spawning), using mechanistic bases. The biogeochemical sub-model successfully simulated phytoplankton dynamics using river nutrient inputs and meteorological factors as forcing functions. Adequate simulation of oyster growth dynamics requires a relevant food quantifier compatible with outputs of the biogeochemical sub-model (i.e. chlorophyll a concentration). We decided to use the phytoplankton carbon concentration as quantifier for food, as it is a better estimator of the energy really available to oysters. The transformation of chlorophyll a concentration into carbon concentration using a variable chlorophyll a to carbon ratio enabled us to improve the simulation of oyster growth especially during the starvation period (i.e. autumn and winter). Once validated, the coupled model was a suitable tool to study the influence of the year-to-year variability of phytoplankton dynamics and water temperature on the gonado-somatic growth of the Pacific oyster. Four years with highly contrasted meteorological conditions (river inputs, water temperature and light) 2000, 2001, 2002 and 2003, were simulated. The years were split into two groups, wet years (2000 and 2001) and dry years (2002 and 2003). Significant variability of the response of oysters to environmental conditions was highlighted between the four scenarios. In the wet years, an increase in loadings of river nutrients and suspended particulate matter led to a shift in the initiation and the magnitude of the phytoplanktonic spring bloom, and consequently to a shift in oyster growth patterns. In contrast, in the dry years, an increase in water temperature—especially during summer—resulted in early spawning. Thus, the gonado-somatic growth pattern of oysters was shown to be sensitive to variations in river loadings and water temperature. In this context, the physiological status of oysters is discussed using a relevant indicator of energy needs.