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Frequent blooms of phytoplankton occur in coastal upwelling zones creating hotspots of biological productivity in the ocean. As cold, nutrient-rich water is brought up to sunlit layers from depth, phytoplankton are also transported upwards to seed surface blooms that are often dominated by diatoms. The physiological response of phytoplankton to this process, commonly referred to as shift-up, is characterized by increases in nitrate assimilation and rapid growth rates. To examine the molecular underpinnings behind this phenomenon, metatranscriptomics was applied to a simulated upwelling experiment using natural phytoplankton communities from the California Upwelling Zone. An increase in diatom growth following 5 days of incubation was attributed to the genera Chaetoceros and Pseudo-nitzschia. Here, we show that certain bloom-forming diatoms exhibit a distinct transcriptional response that coordinates shift-up where diatoms exhibited the greatest transcriptional change following upwelling; however, comparison of co-expressed genes exposed overrepresentation of distinct sets within each of the dominant phytoplankton groups. The analysis revealed that diatoms frontload genes involved in nitrogen assimilation likely in order to outcompete other groups for available nitrogen during upwelling events. We speculate that the evolutionary success of diatoms may be due, in part, to this proactive response to frequently encountered changes in their environment., (© 2018 Society for Applied Microbiology and John Wiley & Sons Ltd.)

Factors that affect the removal of organic carbon by heterotrophic bacterioplankton can impact the rate and magnitude of organic carbon loss in the ocean through the conversion of a portion of consumed organic carbon to CO2. Through enhanced rates of consumption, surface bacterioplankton communities can also reduce the amount of dissolved organic carbon (DOC) available for export from the surface ocean. The present study investigated the direct effects of elevated pCO2 on bacterioplankton removal of several forms of DOC ranging from glucose to complex phytoplankton exudate and lysate, and naturally occurring DOC. Elevated pCO2 (1000-1500 ppm) enhanced both the rate and magnitude of organic carbon removal by bacterioplankton communities compared to low (pre-industrial and ambient) pCO2 (250 -~400 ppm). The increased removal was largely due to enhanced respiration, rather than enhanced production of bacterioplankton biomass. The results suggest that elevated pCO2 can increase DOC consumption and decrease bacterioplankton growth efficiency, ultimately decreasing the amount of DOC available for vertical export and increasing the production of CO2 in the surface ocean.

The summertime North Pacific subtropical gyre has widespread phytoplankton blooms between Hawaii and the subtropical front (∼30°N) that appear as chlorophyll (chl) increases in satellite ocean color data. Nitrogen-fixing diatom symbioses (diatom-diazotroph associations: DDAs) often increase 10(2)-10(3) fold in these blooms and contribute to elevated export flux. In 2008 and 2009, two cruises targeted satellite chlorophyll blooms to examine DDA species abundance, chlorophyll concentration, biogenic silica concentration, and hydrography. Generalized observations that DDA blooms occur when the mixed layer depth is < 70 m are supported, but there is no consistent relationship between mixed layer depth, bloom intensity, or composition; regional blooms between 22-34°N occur within a broader temperature range (21-26°C) than previously reported. In both years, the Hemiaulus-Richelia and Rhizosolenia-Richelia DDAs increased 10(2)-10(3) over background concentrations within satellite-defined bloom features. The two years share a common trend of Hemiaulus dominance of the DDAs and substantial increases in the >10 µm chl a fraction (∼40-90+% of total chl a). Integrated diatom abundance varied 10-fold over <10 km. Biogenic silica concentration tracked diatom abundance, was dominated by the >10 µm size fraction, and increased up to 5-fold in the blooms. The two years differed in the magnitude of the surface chl a increase (2009>2008), the abundance of pennate diatoms within the bloom (2009>2008), and the substantially greater mixed layer depth in 2009. Only the 2009 bloom had sufficient chl a in the >10 µm fraction to produce the observed ocean color chl increase. Blooms had high spatial variability; ocean color images likely average over numerous small events over time and space scales that exceed the individual event scale. Summertime DDA export flux noted at the Hawaii time-series Sta. ALOHA is probably a generalized feature of the eastern N. Pacific north to the subtropical front.

The availability of iron is known to exert a controlling influence on biological productivity in surface waters over large areas of the ocean and may have been an important factor in the variation of the concentration of atmospheric carbon dioxide over glacial cycles. The effect of iron in the Southern Ocean is particularly important because of its large area and abundant nitrate, yet iron-enhanced growth of phytoplankton may be differentially expressed between waters with high silicic acid in the south and low silicic acid in the north, where diatom growth may be limited by both silicic acid and iron. Two mesoscale experiments, designed to investigate the effects of iron enrichment in regions with high and low concentrations of silicic acid, were performed in the Southern Ocean. These experiments demonstrate iron's pivotal role in controlling carbon uptake and regulating atmospheric partial pressure of carbon dioxide.

The North Atlantic is a region of enhanced biogeochemical and climatological importance for the global ocean as it is the site of one of the largest seasonal phytoplankton blooms on the planet. However, there is a lack of understanding of how phytoplankton size influences bloom dynamics and associated nutrient utilization rates, particularly during the decline phase when export to the deep ocean is especially pronounced. Here, we evaluate trends in size‐fractionated carbon, nitrogen, and silicic acid uptake rates in conjunction with environmental parameters to assess these dynamics. In our study, the decline phase of the bloom continued to be highly productive with net primary production (NPP) ranging from 36.4 to 146.6 mmol C m−2 d−1 and approximately 54% of primary production being driven by large phytoplankton cells (≥5 μm) that were primarily utilizing nitrate (mean f‐ratio of 0.77). Entrainment of silicic acid related to deepening of the mixed layer caused by storms increased silicic acid uptake rates to 2.0–5.7 mmol Si m−2 d−1 without concomitant increases in NPP by large cells (silicic acid to carbon uptake ratios averaged 0.12). A companion study in the North Pacific allowed for paired evaluation of these regions. Our results suggest that in highly productive regions where phytoplankton biomass and productivity is distributed across a broad range of cell sizes, such as the North Atlantic, size itself has a stronger influence on nutrient cycling and potential carbon export relative to regions with lower production and a predominance of small (<5 μm) cells, such as the North Pacific. Plain Language Summary: The North Atlantic Ocean experiences a seasonal bloom of phytoplankton. This bloom represents one of the largest removals of anthropogenic carbon to the deep ocean on the planet. Here, we seek to better quantify the mechanisms of this removal by characterizing the amount and composition of phytoplankton present and how much primary production they are engaging in. We sampled during the decline phase of the spring bloom and found the region to be characterized by mostly large phytoplankton primarily utilizing nitrate. When compared to a companion study in the North Pacific, this study confirms the important role cell size and nutrient availability play in determining the degree of productivity in both high and low primary production oceanic regions. Key Points: The decline phase of the North Atlantic spring bloom exhibits transitions in the balance between small and large size‐fractionated primary production dynamicsDuring the bloom decline, diatoms contribute less to primary production but remain a substantial component of phytoplankton biomassPhytoplankton size is a substantial control on ecosystem dynamics in endmember oceanic productivity‐export systems [ABSTRACT FROM AUTHOR]

Coccolithophores are an important group of calcifying marine phytoplankton. Although coccolithophores are not silicified, some species exhibit a requirement for Si in the calcification process. These species also possess a novel protein (SITL) that resembles the SIT family of Si transporters found in diatoms. However, the nature of Si transport in coccolithophores is not yet known, making it difficult to determine the wider role of Si in coccolithophore biology. Here, we show that coccolithophore SITLs act as Na+‐coupled Si transporters when expressed in heterologous systems and exhibit similar characteristics to diatom SITs. We find that CbSITL from Coccolithus braarudii is transcriptionally regulated by Si availability and is expressed in environmental coccolithophore populations. However, the Si requirement of C. braarudii and other coccolithophores is very low, with transport rates of exogenous Si below the level of detection in sensitive assays of Si transport. As coccoliths contain only low levels of Si, we propose that Si acts to support the calcification process, rather than forming a structural component of the coccolith itself. Si is therefore acting as a micronutrient in coccolithophores and natural populations are only likely to experience Si limitation in circumstances where dissolved silicon (DSi) is depleted to extreme levels. [ABSTRACT FROM AUTHOR]

Transitions in phytoplankton community composition are typically attributed to ecological succession even in physically dynamic upwelling systems like the California Current Ecosystem (CCE). An expected succession from a high‐chlorophyll (~ 10 μg L−1) diatom‐dominated assemblage to a low‐chlorophyll (< 1.0 μg L−1) non‐diatom dominated assemblage was observed during a 2013 summer upwelling event in the CCE. Using an interdisciplinary field‐based space‐for‐time approach leveraging both biogeochemical rate measurements and metatranscriptomics, we suggest that this successional pattern was driven primarily by physical processes. An annually recurring mesoscale eddy‐like feature transported significant quantities of high‐phytoplankton‐biomass coastal water offshore. Chlorophyll was diluted during transport, but diatom contributions to phytoplankton biomass and activity (49–62% observed) did not decline to the extent predicted by dilution (18–24% predicted). Under the space‐for‐time assumption, these trends infer diatom biomass and activity and were stimulated during transport. This is hypothesized to result from decreased contact rates with mortality agents (e.g., viruses) and release from nutrient limitation (confirmed by rate data nearshore), as predicted by the Disturbance‐Recovery hypothesis of phytoplankton bloom formation. Thus, the end point taxonomic composition and activity of the phytoplankton assemblage being transported by the eddy‐like feature were driven by physical processes (mixing) affecting physiological (release from nutrient limitation, increased growth) and ecological (reduced mortality) factors that favored the persistence of the nearshore diatoms during transit. The observed connection between high‐diatom‐biomass coastal waters and non‐diatom‐dominated offshore waters supports the proposed mechanisms for this recurring eddy‐like feature moving seed populations of coastal phytoplankton offshore and thereby sustaining their activity. [ABSTRACT FROM AUTHOR]

Abstract: The metabolic activity and growth of phytoplankton taxa drives their ecological function and contribution to biogeochemical processes. We present the first quantitative, taxon‐resolved silica production rates, growth rates, and silica content estimates for co‐occurring diatoms along two cross‐shelf transects off the California coast using the fluorescent tracer 2‐(4‐pyridyl)‐5‐((4‐(2‐dimethylaminoethylaminocarbamoyl)methoxy)phenyl)oxazole, and confocal microscopy. Taxon contribution to total diatom community silica production was predominantly a function of the surface area (SA) of new frustule that each taxon created as opposed to cell abundance or frustule thickness. The influential role of SA made large diatoms disproportionately important to community silica production over short time scales (< 1 d). In some cases, large taxa that comprised only ∼ 15% of numerical cell abundance accounted for over 50% of total community silica production. Over longer time scales relevant to bloom dynamics, the importance of SA declines and growth rate becomes the dominant influence on contribution to production. The relative importance of SA and growth rate in relation to silica production was modeled as the time needed for a smaller, faster‐growing taxon to create more SA than a larger, slower‐growing taxon. Differences in growth rate between the taxa effected the model outcome more than differences in SA. Shifts in relative silica production among taxa are time restricted by finite resources that limit the duration of a bloom. These patterns offer clues as to how taxa respond to their environment and the consequences for both species succession and the potential diatom contribution to elemental cycling. [ABSTRACT FROM AUTHOR]

Six clones of the marine cyanobacterium Synechococcus, representing four major clades, were all found to contain significant amounts of silicon in culture. Growth rate was unaffected by silicic acid, Si( OH)4, concentration between 1 and 120 μM suggesting that Synechococcus lacks an obligate need for silicon (Si). Strains contained two major pools of Si: an aqueous soluble and an aqueous insoluble pool. Soluble pool sizes correspond to estimated intracellular dissolved Si concentrations of 2-24 mM, which would be thermodynamically unstable implying the binding of intracellular soluble Si to organic ligands. The Si content of all clones was inversely related to growth rate and increased with higher [Si( OH)4] in the growth medium. Accumulation rates showed a unique bilinear response to increasing [Si( OH)4] from 1 to 500 μM with the rate of Si acquisition increasing abruptly between 80 and 100 μM Si( OH)4. Although these linear responses imply some form of diffusion-mediated transport, Si uptake rates at low Si (~1 μM Si) were inhibited by orthophosphate, suggesting a role of phosphate transporters in Si acquisition. Theoretical calculations imply that observed Si acquisition rates are too rapid to be supported by lipid-solubility diffusion of Si through the plasmalemma; however, facilitated diffusion involving membrane protein channels may suffice. The data are used to construct a working model of the mechanisms governing the Si content and rate of Si acquisition in Synechococcus. [ABSTRACT FROM AUTHOR]

Factors that affect the removal of organic carbon by heterotrophic bacterioplankton can impact the rate and magnitude of organic carbon loss in the ocean through the conversion of a portion of consumed organic carbon to CO2. Through enhanced rates of consumption, surface bacterioplankton communities can also reduce the amount of dissolved organic carbon (DOC) available for export from the surface ocean. The present study investigated the direct effects of elevated pCO2 on bacterioplankton removal of several forms of DOC ranging from glucose to complex phytoplankton exudate and lysate, and naturally occurring DOC. Elevated pCO2 (1000–1500 ppm) enhanced both the rate and magnitude of organic carbon removal by bacterioplankton communities compared to low (pre-industrial and ambient) pCO2 (250 –~400 ppm). The increased removal was largely due to enhanced respiration, rather than enhanced production of bacterioplankton biomass. The results suggest that elevated pCO2 can increase DOC consumption and decrease bacterioplankton growth efficiency, ultimately decreasing the amount of DOC available for vertical export and increasing the production of CO2 in the surface ocean. [ABSTRACT FROM AUTHOR]

Interactions between dissolved organic carbon (DOC) and bacterioplankton were examined during a diatom and Phaeocystis bloom in the Santa Barbara Channel (SBC) over five days following an upwelling event. The SBC was heterogeneous in physical state (recently upwelled vs. more stratified), nutrient concentration, and productivity, encompassing phytoplankton physiological states from a healthy bloom through the onset of silicon stress. DOC accumulated in the upper 10 m over the bloom, with compositional shifts indicated by chromophoric dissolved organic matter (CDOM) parameters. DOC bioavailability and bacterial growth and community composition responses were assessed with dilution batch-culture bioassays. In these experiments, bacterioplankton DOC usage increased over the bloom, with uptake of 1.5-5.3 μmol L−1 over three days, 1.5-5.7 μmol L−1 over one week, and 1.8-10.8 μmol L−1 over 10 weeks. DOC removal was poorly correlated with traditional proxies of bioavailability (chlorophyll a concentration, elemental ratios of dissolved organic matter, and CDOM). However, bacterial growth efficiency (BGE) was strongly related to in situ conditions, with higher BGEs on fresher, late-bloom DOC. After 10 weeks, 1.6-15.7 μmol L−1 of the DOC that accumulated during the bloom remained unutilized in the bioassays, with higher concentrations of persistent DOC in experiments from senescent bloom physiological states, supporting the putative relationship between phytoplankton blooms and seasonal DOC accumulation in the field. These experiments demonstrate that DOC released by the plankton community during a bloom fuels increased short-term and long-term bacterial activity, enhances presumed trophic transfer via increased BGEs, and leads to the accumulation of persistent, potentially exportable, DOC. © 2015 Association for the Sciences of Limnology and Oceanography [ABSTRACT FROM AUTHOR]

Abstract: The growth of siliceous phytoplankton, mainly diatoms, in the Southern Ocean influences the preformed nutrient inventory in the ocean on a global scale. Silicic acid use by diatoms and deep circulation combine to trap dissolved Si in the Southern Ocean resulting in high levels of silica production and expansive diatom oozes in Southern Ocean sediments. The analysis of the silicon isotope composition of biogenic silica, or opal, and dissolved silicic acid provide insight into the operation of the global marine silicon cycle and the role played by the Southern Ocean in nutrient supply and carbon drawdown, both in the modern and in the past. Silicon isotope studies of diatoms have provided insight into the history of silica production in surface waters, while the analysis of spicules from deep sea sponges has defined both the spatial and the temporal variability of silicic acid concentrations in the water column; together these – and other – proxies reveal variations in the northward flow of Southern Ocean intermediate and mode waters and how changes in Southern Ocean productivity altered their preformed nutrient content. We present a new hypothesis – the “Silicic Acid Ventilation Hypothesis” (SAVH) – to explain the geographical variation of opal-based proxy records, in particular the contrasting patterns of opal burial change found in the low and high latitudes. By understanding the silicon isotope systematics of opal and silicic acid in the modern, we will be able to use opal-based proxies to reconstruct past changes in the Southern Ocean and so investigate its role in global carbon cycling and climate. [Copyright &y& Elsevier]

Abstract: Phytoplankton are a major food resource for filter-feeding organisms occupying intertidal and subtidal habitats of the inner continental shelf. Phytoplankton in these nearshore waters experience different anthropogenic and natural forcing compared to those offshore such that cross-shelf exchange would allow phytoplankton that were produced offshore to serve as a subsidy to inner shelf consumers when phytoplankton production on the shelf is negatively impacted. In the Santa Barbara Channel (SBC) the continental shelf is only a few kilometers wide facilitating exchange with offshore waters. Physical, chemical and biological gradients were examined monthly along a 3-km cross-shelf transect in the SBC from January, 2008 through April, 2009. Chemical and biological distributions followed temporal changes in physical forcing with higher nutrient concentrations and a more intense period of biological production associated with spring upwelling. Chlorophyll was relatively evenly distributed across the shelf during upwelling, but was present at higher concentrations on the inner shelf under stratified conditions. Similarly, cross-shelf gradients in the distribution of dominant phytoplankton genera were weakest during upwelling when blooms of the prymnesiophyte, Phaeocystis, and the diatoms, Eucampia spp. and Thalassiosira spp. occurred across most of the shelf. Upon stratification, blooms were largely confined to the inner shelf within 0.75km of the shoreline with an initial bloom of the diatom Leptocylindrus spp. followed by Pseudo-nitzschia spp. and series of dinoflagellate blooms with Prorocentrum spp. and Lingulodinium spp. attaining the highest abundances. Phytoplankton taxonomic similarity decreased with increasing distance separating stations along the transect and was inversely related to stratification intensity. The observed distribution patterns and the trends in taxonomic similarity imply that for most of the year consumers within rocky intertidal and subtidal ecosystems on the inner shelf are exposed to phytoplankton produced on the continental shelf, and often on the inner shelf. The contribution of phytoplankton produced beyond the shelf break occurs mainly during spring upwelling. [Copyright &y& Elsevier]

Abstract: Silica cycling in the upper 175m of the North Pacific Subtropical Gyre was examined over a two year period (January 2008–December 2009) at the Hawaii Ocean Time-series (HOT) station ALOHA. Silicic acid concentrations in surface waters ranged from 0.6 to 1.6μM, exhibiting no clear seasonal trends. Biogenic silica concentrations and silica production rates increased by an order of magnitude each summer following stratification of the upper 50m reaching values of 157nmolSiL−1 and 81nmolSiL−1 d−1, in 2008 and 2009, respectively. Sea surface height anomalies together with analyses of variability in isothermal surfaces at 150–175m indicated that the summer periods of elevated biogenic silica were associated with anticyclonic mesoscale features during both years. Lithogenic silica concentrations increased in the spring during the known period of maximum atmospheric dust concentrations with maximum values of 36nmolSiL−1 in the upper 10m. Dust deposition would enhance levels of dissolved iron in surface waters, but there was no response of diatom biomass or silica production to increases in near-surface ocean lithogenic silica concentrations suggesting iron sufficiency of diatom silica production rates. Low ambient silicic acid concentrations restricted silica production rates to an average of 43% of maximum potential rates. Si sufficiency only occurred during the summer period when diatom biomass was elevated suggesting that bloom diatoms are adapted to exploit low silicic acid concentrations. Annual silica production at HOT is estimated to be 63mmolSim−2 a−1 with summer blooms contributing 29% of the annual total. Diatoms are estimated to account for 3–7% of total phytoplankton primary productivity, but 9–20% of organic carbon export confirming past suggestions that diatoms are relatively minor contributors to primary productivity and autotrophic biomass, but important contributors to new and export production in oligotrophic open-ocean ecosystems. Annual silica production at HOT is nearly 4-fold lower than estimates at the Bermuda Atlantic Time-series Study (BATS) site in the Sargasso Sea from the 1990s, but annual silica export at the base of the euphotic zone is similar between the two gyres indicating very different balances between silica production and its loss in surface waters. On a relative basis, BATS is a more productive system with respect to silica, where biogenic silica is recycled with high efficiency in surface waters; in contrast the NPSG is a lower productivity system with respect to silica, but where lower recycling efficiency leads to a much higher fraction of new silica production. The two gyres show contrasting long-term trends in diatom biomass as biogenic silica concentrations at HOT have been increasing since 1997, but they have been decreasing at BATS suggesting very different forcing of decadal trends in the contribution of diatoms in carbon cycling between these gyres. Combining the data from both gyres indicates that globally subtropical gyres produce 13TmolSia−1, which is only 51% of previous estimates reducing the contribution of subtropical gyres to 5–7% of global annual marine silica production. [Copyright &y& Elsevier]

Abstract: The relative roles of silicon (Si) and iron (Fe) as limiting nutrients in the eastern equatorial Pacific (EEP) were examined in a series of nine microcosm experiments conducted over two years between 110°W and 140°W longitude. Si and Fe additions had consistently different but synergistic effects on macronutrient use, phytoplankton biomass and phytoplankton community structure. Silicon addition increased silicic acid use and biogenic silica production, but had no significant effect on the use of inorganic nitrogen or orthophosphate, chlorophyll accumulation, particulate inorganic (PIC) carbon accumulation, or plankton community composition relative to controls. That result, together with observations that Si addition increased the cellular Si content of the numerically dominant diatom by ∼50%, indicates that the main effect of Si was to regulate diatom silicification. Like the effect of Si, Fe addition increased the rate of silicic acid use and biogenic silica production and had no effect on PIC production. Unlike the effect of Si, Fe addition also enhanced rates of organic matter production, had no effect on cellular Si content of diatoms, and resulted in the growth of initially rare, large (>40μm) diatoms relative to controls, indicating that Fe limitation acts mainly through its effects on growth rate and phytoplankton community composition. A pennate diatom of the genus Pseudo-nitzschia dominated the diatom assemblage in situ, grew readily in the controls and did not show a strong growth response to either Fe or Si addition suggesting that its growth was regulated by other factors such as grazing or light. Addition of germanium, an inhibitor of diatom cell division, eliminated the effects of Fe on macronutrient use, biogenic silica production and chlorophyll accumulation and phytoplankton community composition, consistent with a predominantly diatom response to Fe addition. The lack of a response of PIC production to Fe suggests that coccolithophores were not Fe limited. Addition of Fe and Si together resulted in the greatest levels of nutrient drawdown and biomass accumulation through the effect of Fe in promoting the growth of large diatoms. The results suggest a form of co-limitation with Si regulating diatom silicification and the rate of biogenic silica production while Fe regulates the production of organic matter through limitation of phytoplankton growth rates, in particular those of large diatoms. The results argue against Si regulation of new production in the EEP under average upwelling conditions. Iron addition was necessary and sufficient to stimulate complete removal of nitrate within the equatorial upwelling zone suggesting that new production was restricted by low ambient dissolved Fe consistent with results from in situ Fe fertilization experiments conducted to the south of the equator outside of the equatorial upwelling zone. [Copyright &y& Elsevier]

Studies the chemical form of dissolved silicon taken up by marine diatoms. Kinetics of silicon use; Use of silicon tracer methodology to determine uptake rates; Variation in half-saturation constant.

Focuses on upward nutrient transport as an important source of new nitrogen to the surface ocean. How the oligotrophic gyres of the open sea are home to a flora that includes the largest known phytoplankton; How these species migrate between deep pools and the surface; How this migration contribute to new production of nitrates.

Phytoplankton exude carbon-rich dissolved organic matter (DOM) upon nutrient stress, yet the ecological role of this exudate in stimulating bacterial growth is not well understood. We harvested DOM produced by four coastal diatoms ( Skeletonema marinoi , Chaetoceros socialis , Thalassiosira weissflogii , and Odontella aurita ) subjected to depletion of nitrogen (N), silicon (Si), or N + Si simultaneously in batch culture and assessed its bioavailability to natural bacterioplankton communities using dilution batch-culture remineralization bioassays. Short-term, ecologically-relevant responses were affected both by diatom source species and by the nutrient stress under which the DOM was produced. Si-stress DOM was generally more bioavailable over the first week than that produced under N or N + Si stress, and led to higher bacterial growth efficiencies. In contrast, the amount of diatom-derived total organic carbon (TOC) that persisted over months in the same experiments differed among source diatom species, with no evidence of a nutrient stress effect. These results nuance the carbon overflow hypothesis of DOM release, which has been suggested to be maladaptive due to the possibility of DOC release allowing bacterioplankton to better compete for the same nutrient that is limiting the phytoplankton. Our results suggest that bacterial activity is most promoted when diatoms are limited by Si, a nutrient that is not subject to competition with bacteria. They imply that the identity of the nutrient that terminates a diatom bloom impacts heterotrophic activity in the short term; however, source diatom species has the greater influence on DOC persistence in surface waters and its potential export from the system. [ABSTRACT FROM AUTHOR]

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 (fNC), 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. [Copyright &y& Elsevier]