Global Marine Primary Production Research Articles

Environmental Regulation of Photosynthetically Produced Dissolved Organic Carbon by Phytoplankton Along a Subtropical Estuarine Bay

Photosynthetically produced dissolved organic carbon (PDOC) released by marine phytoplankton has an important significance on the marine carbon cycle and on the growth of heterotopic bacteria (HB). However, PDOC is often neglected in the estimation of global marine primary productivity and carbon sequestration capacity. This work studied the issue of PDOC and its environmental regulation mechanism through field investigation and lab experiments in an estuarine bay during southwest and northeast monsoon. We used the percentage of extracellular release (PER) as a key indicator to evaluate the contribution of PDOC to total primary productivity (TPP). We also compared PER among different seasons and sectors and then analyzed the inter-relationship between PDOC and bacterial carbon demand (BCD), size-fractionated phytoplankton, bacterial production (BP), and TPP. We finally discussed the impact factors of PDOC productivity. The results showed that the average contribution of PDOC to TPP in Qinzhou bay could reach 15% during two seasons, which satisfies about 25% of the carbon requirement by HB in the bay. Multiple factors contribute to the seasonal (SW monsoon 13% < NE monsoon 18%) and sectoral variation (outer bay 26% > middle bay 17% > inner bay 7%) in PER, the most significant of which are salinity and nitrogen to phosphorus ratios. PER is also related to phytoplankton community structure and nutrient limitation, the higher PER in the outer bay is attributed to the dual effect of picophytoplankton being the predominant species and a severe imbalance in the nitrogen to phosphorus ratio, both of which lead to increased phytoplankton PDOC release. On the other hand, the lower PER in the inner bay was mainly due to the dominance of microphytoplankton and nitrogen to phosphorus ratio which is close to the redfield ratio. There is still a non-uniform conclusion on the relationship between PER and TPP, suggesting that this relationship needs to be evaluated not only in a horizontal-independent way but also through seasonal and vertical dimensions. The non-linear relationship between PDOC and BCD found in this study potentially indicated weak coupling between phytoplankton exudation and bacterial metabolism. In addition to phytoplankton exudate, HB is still dependent on other sources of DOC to meet their carbon requirements.

Spatio-Temporal Variation of Synechococcus Assemblages at DNA and cDNA Levels in the Tropical Estuarine and Coastal Waters.

Synechococcus is a major contributor to global marine primary production. Here, its spatio-temporal variations in abundance and phylogenetic structure were studied at three stations of the South China Sea at both DNA and cDNA levels. Synechococcus cell abundance was lowest in March, but highest in October at two coastal stations. Its abundance was higher at the estuarine station, which reached a peak value of 1.36 × 105 cells/ml in April, owing to the nitrogen nutrients discharged from the Sanya River. Gene and gene transcript abundances of four Synechococcus lineages, clades II, III, VIII, and S5.3, were studied by quantitative PCR, which showed that clade II was the most abundant lineage at both DNA and cDNA levels. High-throughput sequencing revealed that, at the DNA level, Synechococcus assemblage was dominated by clade SY4 (a novel clade defined in this study), S5.2, and clade II in the coastal waters and was dominated by freshwater/S5.2 Synechococcus, reaching a value up to 88.61% in June, in estuarine waters. Changes in salinity and nutrient concentration caused by seasonal monsoonal forcing and river discharge were the key determinants of the spatio-temporal variation in Synechococcus assemblages at the DNA level. In comparison, high dissimilation among samples at the same stations and in the same seasons leads to the imperceptible spatio-temporal variation pattern of Synechococcus assemblages at the cDNA level. Furthermore, co-occurrence networks disclosed that Synechococcus community had closer and more complex internal interactions at the cDNA level. These discrepancies highlighted the necessity to study Synechococcus assemblages at both DNA and cDNA levels.

Riverine nitrogen supply to the global ocean and its limited impact on global marine primary production: a feedback study using an Earth system model

Abstract. A common notion is that negative feedbacks stabilize the natural marine nitrogen inventory. Recent modeling studies have shown, however, some potential for localized positive feedbacks leading to substantial nitrogen losses in regions where nitrogen fixation and denitrification occur in proximity to each other. Here we include dissolved nitrogen from river discharge in a global 3-D ocean biogeochemistry model and study the effects on near-coastal and remote-open-ocean biogeochemistry. We find that at a steady state the biogeochemical feedbacks in the marine nitrogen cycle, nitrogen input from biological N2 fixation, and nitrogen loss via denitrification mostly compensate for the imposed yearly addition of 22.8 to 45.6 Tg of riverine nitrogen and limit the impact on global marine productivity to < 2 %. Global experiments that regionally isolate river nutrient input show that the sign and strength of the feedbacks depend on the location of the river discharge and the oxygen status of the receiving marine environment. Marine productivity generally increases in proximity to the nitrogen input, but we also find a decline in productivity in the modeled Bay of Bengal and near the mouth of the Amazon River. While most of the changes are located in shelf and near-coastal oceans, nitrogen supply from the rivers can impact the open ocean, due to feedbacks or knock-on effects.

Meta-analysis of multiple driver effects on marine phytoplankton highlights modulating role of pCO2.

Responses of marine primary production to a changing climate are determined by a concert of multiple environmental changes, for example in temperature, light, pCO2 , nutrients, and grazing. To make robust projections of future global marine primary production, it is crucial to understand multiple driver effects on phytoplankton. This meta-analysis quantifies individual and interactive effects of dual driver combinations on marine phytoplankton growth rates. Almost 50% of the single-species laboratory studies were excluded because central data and metadata (growth rates, carbonate system, experimental treatments) were insufficiently reported. The remaining data (42 studies) allowed for the analysis of interactions of pCO2 with temperature, light, and nutrients, respectively. Growth rates mostly respond non-additively, whereby the interaction with increased pCO2 profusely dampens growth-enhancing effects of high temperature and high light. Multiple and single driver effects on coccolithophores differ from other phytoplankton groups, especially in their high sensitivity to increasing pCO2 . Polar species decrease their growth rate in response to high pCO2 , while temperate and tropical species benefit under these conditions. Based on the observed interactions and projected changes, we anticipate primary productivity to: (a) first increase but eventually decrease in the Arctic Ocean once nutrient limitation outweighs the benefits of higher light availability; (b) decrease in the tropics and mid-latitudes due to intensifying nutrient limitation, possibly amplified by elevated pCO2 ; and (c) increase in the Southern Ocean in view of higher nutrient availability and synergistic interaction with increasing pCO2 . Growth-enhancing effect of high light and warming to coccolithophores, mainly Emiliania huxleyi, might increase their relative abundance as long as not offset by acidification. Dinoflagellates are expected to increase their relative abundance due to their positive growth response to increasing pCO2 and light levels. Our analysis reveals gaps in the knowledge on multiple driver responses and provides recommendations for future work on phytoplankton.

Genome Sequences of Synechococcus sp. Strain MIT S9220 and Cocultured Cyanophage SynMITS9220M01.

Synechococcus bacteria are unicellular cyanobacteria that contribute significantly to global marine primary production. We report the nearly complete genome sequence of Synechococcus sp. strain MIT S9220, which lacks the nitrate utilization genes present in most marine Synechococcus genomes. Assembly also produced the complete genome sequence of a cyanophage present in the MIT S9220 culture.

Elevated sedimentary removal of Fe, Mn, and trace elements following a transient oxygenation event in the Eastern Gotland Basin, central Baltic Sea

Iron, manganese, and trace elements play an important role in the marine carbon cycle as they are limiting nutrients for marine primary productivity. Water column concentrations of these bio-essential elements are controlled by the balance between input and removal, with burial in marine sediments being the main sink. The efficiency of this burial sink is dependent on the redox state of the water column, with sediments underlying a sulphidic (euxinic) water column being the most efficient sinks for Fe, but also Mn and trace elements (Co, Cd, Ni, Mo, As, W, V, and U). Transient changes in ocean redox state can hence affect trace element burial, and correspondingly, the ocean’s trace element inventory, but the impact of transient oxygenation events on trace element cycling is currently not well understood.Here, we investigate the impact of a natural oxygenation event on trace element release and burial in sediments of the Eastern Gotland Basin (EGB), a sub-basin of the Baltic Sea. After being anoxic (<0.5 µM O2) for ∼10 years, the deep waters of the EGB experienced a natural oxygenation event (Major Baltic Inflow, MBI) in 2015. Following this oxygenation event, we deployed benthic chamber landers along a depth transect in the EGB in April 2016, 2017 and 2018. We complemented these in situ flux measurements with analyses of water column, solid phase and pore water chemistry. Overall, the event increased the benthic effluxes of dissolved trace elements, though particular responses were element-specific and were caused by different mechanisms. Enhanced fluxes of Cd and U were caused by oxidative remobilisation, while Ni showed little response to the inflow of oxygen. In contrast, enhanced release of Co, Mo, As, W, and V was caused by the enhanced transient input of Mn oxides into the sediment, whereas Fe oxides were of minor importance. Following the dissolution of the oxides in the sediment, Mn and W were nearly completely recycled back to the water column, while fractions of Fe, Co, Mo, As, and V were retained in the sediment. Our results suggest that transient oxygenation events in euxinic basins may decrease the water column inventory of certain trace elements (Fe, Co, Mo, As, and V), thus potentially affecting global marine primary productivity on longer timescales.

Consistent trophic amplification of marine biomass declines under climate change.

The impact of climate change on the marine food web is highly uncertain. Nonetheless, there is growing consensus that global marine primary production will decline in response to future climate change, largely due to increased stratification reducing the supply of nutrients to the upper ocean. Evidence to date suggests a potential amplification of this response throughout the trophic food web, with more dramatic responses at higher trophic levels. Here we show that trophic amplification of marine biomass declines is a consistent feature of the Coupled Model Intercomparison Project Phase 5 (CMIP5) Earth System Models, across different scenarios of future climate change. Under the business-as-usual Representative Concentration Pathway 8.5 (RCP8.5) global mean phytoplankton biomass is projected to decline by 6.1%±2.5% over the twenty-first century, while zooplankton biomass declines by 13.6%±3.0%. All models project greater relative declines in zooplankton than phytoplankton, with annual zooplankton biomass anomalies 2.24±1.03 times those of phytoplankton. The low latitude oceans drive the projected trophic amplification of biomass declines, with models exhibiting variable trophic interactions in the mid-to-high latitudes and similar relative changes in phytoplankton and zooplankton biomass. Under the assumption that zooplankton biomass is prey limited, an analytical explanation of the trophic amplification that occurs in the low latitudes can be derived from generic plankton differential equations. Using an ocean biogeochemical model, we show that the inclusion of variable C:N:P phytoplankton stoichiometry can substantially increase the trophic amplification of biomass declines in low latitude regions. This additional trophic amplification is driven by enhanced nutrient limitation decreasing phytoplankton N and P content relative to C, hence reducing zooplankton growth efficiency. Given that most current Earth System Models assume that phytoplankton C:N:P stoichiometry is constant, such models are likely to underestimate the extent of negative trophic amplification under projected climate change.

Photosynthesis–irradiance parameters of marine phytoplankton: synthesis of a global data set

Abstract. The photosynthetic performance of marine phytoplankton varies in response to a variety of factors, environmental and taxonomic. One of the aims of the MArine primary Production: model Parameters from Space (MAPPS) project of the European Space Agency is to assemble a global database of photosynthesis–irradiance (P-E) parameters from a range of oceanographic regimes as an aid to examining the basin-scale variability in the photophysiological response of marine phytoplankton and to use this information to improve the assignment of P-E parameters in the estimation of global marine primary production using satellite data. The MAPPS P-E database, which consists of over 5000 P-E experiments, provides information on the spatio-temporal variability in the two P-E parameters (the assimilation number, PmB, and the initial slope, αB, where the superscripts B indicate normalisation to concentration of chlorophyll) that are fundamental inputs for models (satellite-based and otherwise) of marine primary production that use chlorophyll as the state variable. Quality-control measures consisted of removing samples with abnormally high parameter values and flags were added to denote whether the spectral quality of the incubator lamp was used to calculate a broad-band value of αB. The MAPPS database provides a photophysiological data set that is unprecedented in number of observations and in spatial coverage. The database will be useful to a variety of research communities, including marine ecologists, biogeochemical modellers, remote-sensing scientists and algal physiologists. The compiled data are available at https://doi.org/10.1594/PANGAEA.874087 (Bouman et al., 2017).

Emergent constraints on projections of declining primary production in the tropical oceans

Marine primary production is a fundamental component of the Earth system, providing the main source of food and energy to the marine food web, and influencing the concentration of atmospheric CO$_2$ (refs. 1,2). Earth system model (ESM) projections of global marine primary production are highly uncertain with models projecting both increases (refs. 3,4) and declines of up to 20% by 2100 (refs. 5,6) This uncertainty is predominantly driven by the sensitivity of tropical ocean primary production to climate change, with the latest ESMs suggesting twenty-first-century tropical declines of between 1 and 30% (refs 5,6). Here we identify an emergent relationship (refs. 7-11) between the long-term sensitivity of tropical ocean primary production to rising equatorial zone sea surface temperature (SST) and the interannual sensitivity of primary production to El Nino/Southern Oscillation (ENSO)-driven SST anomalies. Satellite-based observations of the ENSO sensitivity of tropical primary production are then used to constrain projections of the long-term climate impact on primary production. We estimate that tropical primary production will decline by 3 ± 1% per kelvin increase in equatorial zone SST. Under a business-as-usual emissions scenario this results in an 11 ± 6% decline in tropical marine primary production and a 6 ± 3% decline in global marine primary production by 2100.

Phylogeography and pigment type diversity of Synechococcus cyanobacteria in surface waters of the northwestern pacific ocean.

The widespread unicellular cyanobacteria Synechococcus are major contributors to global marine primary production. Here, we report their abundance, phylogenetic diversity (as assessed using the RNA polymerase gamma subunit gene rpoC1) and pigment diversity (as indirectly assessed using the laterally transferred cpeBA genes, encoding phycoerythrin-I) in surface waters of the northwestern Pacific Ocean, sampled over nine distinct cruises (2008-2015). Abundance of Synechococcus was low in the subarctic ocean and South China Sea, intermediate in the western subtropical Pacific Ocean, and the highest in the Japan and East China seas. Clades I and II were by far the most abundant Synechococcus lineages, the former dominating in temperate cold waters and the latter in (sub)tropical waters. Clades III and VI were also fairly abundant in warm waters, but with a narrower distribution than clade II. One type of chromatic acclimater (3dA) largely dominated the Synechococcus communities in the subarctic ocean, while another (3dB) and/or cells with a fixed high phycourobilin to phycoerythrobilin ratio (pigment type 3c) predominated at mid and low latitudes. Altogether, our results suggest that the variety of pigment content found in most Synechococcus clades considerably extends the niches that they can colonize and therefore the whole genus habitat.

Dissolved organic carbon fluxes in the Middle Atlantic Bight: An integrated approach based on satellite data and ocean model products.

Continental margins play an important role in global carbon cycle, accounting for 15-21% of the global marine primary production. Since carbon fluxes across continental margins from land to the open ocean are not well constrained, we undertook a study to develop satellite algorithms to retrieve dissolved organic carbon (DOC) and combined these satellite data with physical circulation model products to quantify the shelf boundary fluxes of DOC for the U.S. Middle Atlantic Bight (MAB). Satellite DOC was computed through seasonal relationships of DOC with colored dissolved organic matter absorption coefficients, which were derived from an extensive set of in situ measurements. The multiyear time series of satellite-derived DOC stocks (4.9 Teragrams C; Tg) shows that freshwater discharge influences the magnitude and seasonal variability of DOC on the continental shelf. For the 2010-2012 period studied, the average total estuarine export of DOC into the MAB shelf is 0.77 Tg C yr-1 (year). The integrated DOC tracer fluxes across the shelf boundaries are 12.1 Tg C yr-1 entering the MAB from the southwest alongshore boundary, 18.5 Tg C yr-1 entering the MAB from the northeast alongshore boundary, and 29.0 Tg C yr-1 flowing out of the MAB across the entire length of the 100 m isobath. The magnitude of the cross-shelf DOC flux is quite variable in time (monthly) and space (north to south). The highly dynamic exchange of water along the shelf boundaries regulates the DOC budget of the MAB at subseasonal time scales.

Drivers and uncertainties of future global marine primary production in marine ecosystem models

Abstract. Past model studies have projected a global decrease in marine net primary production (NPP) over the 21st century, but these studies focused on the multi-model mean rather than on the large inter-model differences. Here, we analyze model-simulated changes in NPP for the 21st century under IPCC's high-emission scenario RCP8.5. We use a suite of nine coupled carbon–climate Earth system models with embedded marine ecosystem models and focus on the spread between the different models and the underlying reasons. Globally, NPP decreases in five out of the nine models over the course of the 21st century, while three show no significant trend and one even simulates an increase. The largest model spread occurs in the low latitudes (between 30° S and 30° N), with individual models simulating relative changes between −25 and +40 %. Of the seven models diagnosing a net decrease in NPP in the low latitudes, only three simulate this to be a consequence of the classical interpretation, i.e., a stronger nutrient limitation due to increased stratification leading to reduced phytoplankton growth. In the other four, warming-induced increases in phytoplankton growth outbalance the stronger nutrient limitation. However, temperature-driven increases in grazing and other loss processes cause a net decrease in phytoplankton biomass and reduce NPP despite higher growth rates. One model projects a strong increase in NPP in the low latitudes, caused by an intensification of the microbial loop, while NPP in the remaining model changes by less than 0.5 %. While models consistently project increases NPP in the Southern Ocean, the regional inter-model range is also very substantial. In most models, this increase in NPP is driven by temperature, but it is also modulated by changes in light, macronutrients and iron as well as grazing. Overall, current projections of future changes in global marine NPP are subject to large uncertainties and necessitate a dedicated and sustained effort to improve the models and the concepts and data that guide their development.

Satellite remote sensing of ultraviolet irradiance on the ocean surface

Ultraviolet (UV) radiation has a significant influence on marine biological processes and primary productivity; however, the existing ocean color satellite sensors seldom contain UV bands. A look-up table of wavelength-integrated UV irradiance (280–400 nm) on the sea surface is established using the coupled ocean atmosphere radiative transfer (COART) model. On the basis of the look-up table, the distributions of the UV irradiance at middle and low latitudes are inversed by using the satellite-derived atmospheric products from the Aqua satellite, including aerosol optical thickness at 550 nm, ozone content, liquid water path, and the total precipitable water. The validation results show that the mean relative difference of the 10 d rolling averaged UV irradiance between the satellite retrieval and field observations is 8.20% at the time of satellite passing and 13.95% for the daily dose of UV. The monthly-averaged UV irradiance and daily dose of UV retrieved by satellite data show a good correlation with the in situ data, with mean relative differences of 6.87% and 8.43%, respectively. The sensitivity analysis of satellite inputs is conducted. The liquid water path representing the condition of cloud has the highest effect on the retrieval of the UV irradiance, while ozone and aerosol have relatively lesser effect. The influence of the total precipitable water is not significant. On the basis of the satellite-derived UV irradiance on the sea surface, a preliminary simple estimation of ultraviolet radiation’s effects on the global marine primary productivity is presented, and the results reveal that ultraviolet radiation has a non-negligible effect on the estimation of the marine primary productivity.

Drivers and uncertainties of future global marine primary production in marine ecosystem models

Drivers and uncertainties of future global marine primary production in marine ecosystem models

Modeling the global cycle of marine dissolved organic matter and its influence on marine productivity

A three-dimensional numerical model is developed for studies on the marine biogeochemical cycles by dealing with coupling of planktonic and microbial processes. The coupling is achieved by explicitly representing generation and consumption of dissolved organic matter (DOM). The model is applied to investigate mechanisms by which the global cycle of marine DOM influences marine productivity. Two categories, biodegradable (DOMb) and refractory (DOMr), are considered for DOM. DOMb is generated through phytoplankton exudation, zooplankton excretion, detritus decomposition, and photodissociation of DOMr. Bacteria consume DOMb and generate DOMr. After the long-term adjustment, the observed amount of DOM was reproduced from homogenous conditions of nutrients and plankton. Spatial distribution of bacterial abundance is reasonably validated against recently observed large-scale data. Sensitivity experiments indicated that: (i) over a multi-decadal timescale, contribution of DOMr to global biogeochemical cycles is negligible, (ii) the existence of DOM significantly reduces the global marine primary productivity, and (iii) DOMb originated from phytoplankton exudation is transported to subtropical regions, leading to enhanced primary productivity there as a consequence of nutrient supply associated with remineralization of transported DOMb.

Effects of growth and dissolution on the fractionation of silicon isotopes by estuarine diatoms

Studies of silicon (Si) isotope fractionation during diatom growth in open ocean systems have documented lower Si isotopic values (δ30Si) in the biogenic silica of diatom frustules compared to dissolved silicon. Recent findings also indicate that Si isotope fractionation occurs during dissolution of diatom frustules, producing higher δ30Si values in the remaining biogenic silica. This study focuses on diatoms from high production areas in estuarine and coastal areas that represent approximately 30–50% of the global marine primary production. Two species of diatoms, Thalassiosira baltica and Skeletonema marinoi, were isolated from the brackish Baltic Sea, one of the largest estuarine systems in the world. These species were used for laboratory investigations of Si isotope fractionation during diatom growth and the subsequent dissolution of the diatom frustules. Both species of diatoms give an identical Si isotope fractionation factor during growth of −1.50±0.36‰ (2σ) for 30Si, which falls in the range of −2.09‰ to −0.55‰ of published data. Our results also suggest a dissolution-induced Si isotope fractionation factor of −0.86‰ at early stage of dissolution, but this effect was observed only in DSi and no significant Si isotope change was observed for BSi. The growth and dissolution results are applied to a Baltic Sea sediment core to reconstruct DSi utilization by diatoms, and found to be in agreement with the observed DSi uptake rates in the overlying water column during diatom growth.

NW European shelf under climate warming: implications for open ocean – shelf exchange, primary production, and carbon absorption

Abstract. Shelves have been estimated to account for more than one-fifth of the global marine primary production. It has been also conjectured that shelves strongly influence the oceanic absorption of anthropogenic CO2 (carbon shelf pump). Owing to their coarse resolution, currently applied global climate models are inappropriate to investigate the impact of climate change on shelves and regional models do not account for the complex interaction with the adjacent open ocean. In this study, a global ocean general circulation model and biogeochemistry model were set up with a distorted grid providing a maximal resolution for the NW European shelf and the adjacent northeast Atlantic. Using model climate projections we found that already a~moderate warming of about 2.0 K of the sea surface is linked with a reduction by ~ 30% of the biological production on the NW European shelf. If we consider the decline of anthropogenic riverine eutrophication since the 1990s, the reduction of biological production amounts is even larger. The relative decline of NW European shelf productivity is twice as strong as the decline in the open ocean (~ 15%). The underlying mechanism is a spatially well confined stratification feedback along the continental shelf break. This feedback reduces the nutrient supply from the deep Atlantic to about 50%. In turn, the reduced productivity draws down CO2 absorption in the North Sea by ~ 34% at the end of the 21st century compared to the end of the 20th century implying a strong weakening of shelf carbon pumping. Sensitivity experiments with diagnostic tracers indicate that not more than 20% of the carbon absorbed in the North Sea contributes to the long-term carbon uptake of the world ocean. The rest remains within the ocean's mixed layer where it is exposed to the atmosphere. The predicted decline in biological productivity, and decrease of phytoplankton concentration (in the North Sea by averaged 25%) due to reduced nutrient imports from the deeper Atlantic will probably affect the local fish stock negatively and therefore fisheries in the North Sea.

From the chlorophyll &amp;lt;i&amp;gt;a&amp;lt;/i&amp;gt; in the surface layer to its vertical profile: a Greenland Sea relationship for satellite applications

Abstract. Current estimates of global marine primary production range over a factor of two. Improving these estimates requires an accurate knowledge of the chlorophyll vertical profiles, since they are the basis for most primary production models. At high latitudes, the uncertainty in primary production estimates is larger than globally, because here phytoplankton absorption shows specific characteristics due to the low-light adaptation, and in situ data and ocean colour observations are scarce. To date, studies describing the typical chlorophyll profile based on the chlorophyll in the surface layer have not included the Arctic region, or, if it was included, the dependence of the profile shape on surface concentration was neglected. The goal of our study was to derive and describe the typical Greenland Sea chlorophyll profiles, categorized according to the chlorophyll concentration in the surface layer and further monthly resolved profiles. The Greenland Sea was chosen because it is known to be one of the most productive regions of the Arctic and is among the regions in the Arctic where most chlorophyll field data are available. Our database contained 1199 chlorophyll profiles from R/Vs Polarstern and Maria S. Merian cruises combined with data from the ARCSS-PP database (Arctic primary production in situ database) for the years 1957–2010. The profiles were categorized according to their mean concentration in the surface layer, and then monthly median profiles within each category were calculated. The category with the surface layer chlorophyll (CHL) exceeding 0.7 mg C m−3 showed values gradually decreasing from April to August. A similar seasonal pattern was observed when monthly profiles were averaged over all the surface CHL concentrations. The maxima of all chlorophyll profiles moved from the greater depths to the surface from spring to late summer respectively. The profiles with the smallest surface values always showed a subsurface chlorophyll maximum with its median magnitude reaching up to three times the surface concentration. While the variability of the Greenland Sea season in April, May and June followed the global non-monthly resolved relationship of the chlorophyll profile to surface chlorophyll concentrations described by the model of Morel and Berthon (1989), it deviated significantly from the model in the other months (July–September), when the maxima of the chlorophyll are at quite different depths. The Greenland Sea dimensionless monthly median profiles intersected roughly at one common depth within each category. By applying a Gaussian fit with 0.1 mg C m−3 surface chlorophyll steps to the median monthly resolved chlorophyll profiles of the defined categories, mathematical approximations were determined. They generally reproduce the magnitude and position of the CHL maximum, resulting in an average 4% underestimation in Ctot (and 2% in rough primary production estimates) when compared to in situ estimates. These mathematical approximations can be used as the input to the satellite-based primary production models that estimate primary production in the Arctic regions.

Multiparametric Analyses Reveal the pH-Dependence of Silicon Biomineralization in Diatoms

Diatoms, the major contributors of the global biogenic silica cycle in modern oceans, account for about 40% of global marine primary productivity. They are an important component of the biological pump in the ocean, and their assemblage can be used as useful climate proxies; it is therefore critical to better understand the changes induced by environmental pH on their physiology, silicification capability and morphology. Here, we show that external pH influences cell growth of the ubiquitous diatom Thalassiosira weissflogii, and modifies intracellular silicic acid and biogenic silica contents per cell. Measurements at the single-cell level reveal that extracellular pH modifications lead to intracellular acidosis. To further understand how variations of the acid-base balance affect silicon metabolism and theca formation, we developed novel imaging techniques to measure the dynamics of valve formation. We demonstrate that the kinetics of valve morphogenesis, at least in the early stages, depends on pH. Analytical modeling results suggest that acidic conditions alter the dynamics of the expansion of the vesicles within which silica polymerization occurs, and probably its internal pH. Morphological analysis of valve patterns reveals that acidification also reduces the dimension of the nanometric pores present on the valves, and concurrently overall valve porosity. Variations in the valve silica network seem to be more correlated to the dynamics and the regulation of the morphogenesis process than the silicon incorporation rate. These multiparametric analyses from single-cell to cell-population levels demonstrate that several higher-level processes are sensitive to the acid-base balance in diatoms, and its regulation is a key factor for the control of pattern formation and silicon metabolism.

Processes controlling the Si-isotopic composition in the Southern Ocean and application for paleoceanography

Abstract. Southern Ocean biogeochemical processes have an impact on global marine primary production and global elemental cycling, e.g. by likely controlling glacial-interglacial pCO2 variation. In this context, the natural silicon isotopic composition (δ30Si) of sedimentary biogenic silica has been used to reconstruct past Si-consumption:supply ratios in the surface waters. We present a new dataset in the Southern Ocean from a IPY-GEOTRACES transect (Bonus-GoodHope) which includes for the first time summer δ30Si signatures of suspended biogenic silica (i) for the whole water column at three stations and (ii) in the mixed layer at seven stations from the subtropical zone up to the Weddell Gyre. In general, the isotopic composition of biogenic opal exported to depth was comparable to the opal leaving the mixed layer and did not seem to be affected by any diagenetic processes during settling, even if an effect of biogenic silica dissolution cannot be ruled out in the northern part of the Weddell Gyre. We develop a mechanistic understanding of the processes involved in the modern Si-isotopic balance, by implementing a mixed layer model. We observe that the accumulated biogenic silica (sensu Rayleigh distillation) should satisfactorily describe the δ30Si composition of biogenic silica exported out of the mixed layer, within the limit of the current analytical precision on the δ30Si. The failures of previous models (Rayleigh and steady state) become apparent especially at the end of the productive period in the mixed layer, when biogenic silica production and export are low. This results from (1) a higher biogenic silica dissolution:production ratio imposing a lower net fractionation factor and (2) a higher Si-supply:Si-uptake ratio supplying light Si-isotopes into the mixed layer. The latter effect is especially expressed when the summer mixed layer becomes strongly Si-depleted, together with a large vertical silicic acid gradient, e.g. in the Polar Front Zone and at the Polar Front.