Phytoplankton Exudates Research Articles

Biogeochemical factors regulating photosynthetically dissolved organic carbon produced by phytoplankton in the Taiwan Strait

The distribution and mechanisms of photosynthetically dissolved organic carbon (PDOC) released by marine phytoplankton are frequently neglected and inadequately understood because most studies on carbon sequestration capacity have focused on photosynthetic particulate organic carbon. In this study, percentage extracellular release (PER) and its environmental influencing factors were investigated for 10 cruises in the Taiwan Strait during 2006–2023. The results indicated that the PER increased horizontally from the nearshore to the off-shelf and vertically from the surface to the bottom within the euphotic zone. PER tends to be low in eutrophic waters such as upwellings and estuaries and high in oligotrophic waters. The study revealed that the average contribution of PDOC to total primary productivity (TPP) in the Taiwan Strait could reach 18.2 ± 11.7%, which is similar to the previously estimated global oceanic values. PDOC production satisfied approximately 25% the carbon requirements of heterotropic bacteria (HB). A detailed analysis of the PER combined with model simulations proved that the distribution of the PER in the Taiwan Strait was caused by the joint contribution of irradiance, size-fractionated phytoplankton, and nutrient stoichiometry. Our results contradict the view that the PER is a constant factor that is unaffected by TPP. However, there was a significant negative correlation between the PER and TPP. The PDOC was always lower than the bacterial carbon demand for a broad range of bacterial growth efficiencies, suggesting a weak coupling between phytoplankton exudation and bacterial metabolism. This challenges the idea that there is a well-coupled relationship between bacteria and phytoplankton present on the continental shelf. These findings indicate significant discrepancies in PDOC mechanisms and the quantitative importance of nearshore eutrophic and off-shelf oligotrophic environments. Consequently, it is unwise to use uniform PERs without differentiation under trophic conditions when reevaluating and appraising marine carbon fixation.

A first estimation of the role of penguin guano on copper cycling and organic speciation in Antarctic coastal waters

Cu is a vital micronutrient, but free Cu ions (Cu2+) in seawater, even at subnanomolar concentrations, can impede phytoplankton growth. Natural Cu complexation with organic ligands regulates Cu acquisition and, in most instances, reduces Cu2+ concentrations below toxic thresholds. Along the Antarctic coast, the sources and sinks of Cu and its associated ligands remain poorly defined. Despite the high productivity in the area, there are no studies on the role of trophic transfer in Cu cycling.This study explores penguin guano release of Cu and Cu ligands and its potential in neutralizing copper toxicity along the Antarctic coast. We collected guano in a Chinstrap penguin nesting location in the West coast of Deception Island and extracted its components into aqueous solution imitating natural processes. Copper concentration in guano was 0.4 mg (dry weight g)−1 constituting a potential toxic threat and showed biomagnification with respect to krill. Surface seawater samples collected from various locations varying in penguin activity, were analyzed to assess the potential influence of guano on the area. Visual examination and elevated levels of Al suggested that a substantial portion of guano was lithogenic. Consequently, only a modest 16 % of the total Cu present in guano could be extracted using mechanical methods. Notably, the concentrations of the extracted organic ligands were approximately 23 times higher than the concentrations of the extracted Cu. This significant presence of ligands effectively nullifies any potential toxicity that could have arisen from free Cu2+ ions. Guano ligands' conditional stability constants were lower than those in surface seawater suggesting phytoplankton exudation was the main ligand source in the area. Overall, guano acts as a key node for Cu cycling in coastal Antarctic waters but its deleterious potential is neutralized by ligands from krill digestion and the high background concentration of phytoplankton exudates.

Fluorescent properties of marine phytoplankton exudates and lability to marine heterotrophic prokaryotes degradation

AbstractExudates by the diatom Phaeodactylum tricornutum were incubated with a natural community of marine heterotrophic prokaryotes for 24 d in order to investigate the link between the biological lability and the molecular weight, fluorescence, and polarity of phytoplankton dissolved organic matter (DOM). Dissolved organic carbon (DOC) removal, changes in fluorescence and in the heterotrophic prokaryote abundance were followed over time both in the total exudates and in the low‐ and high‐molecular‐weight fractions. To detect changes in the polarity of proteins, reverse‐phase high‐performance liquid chromatography (HPLC) was applied to the high‐molecular‐weight fraction. Our results indicate that freshly produced phytoplankton DOM exhibits a dynamic pattern of degradation that is accompanied by large changes in the growth efficiency of the bacterial community that are likely related to changes in DOM quality. Approximately 20% of high‐molecular‐weight DOM and 40% of fluorescence attributed to protein‐like DOM were degraded over the first days of the incubation indicating that protein‐like DOM is likely a labile component of phytoplankton exudates. In contrast, fluorescence measurements suggest that humic‐like substances are resistant to bacterial degradation over the 24 d of the experiment. Despite fluctuations in the short‐term rates of high‐molecular‐weight and low‐molecular‐weight DOM removal, the relative contributions of these fractions to DOM pool were similar in the fresh exudates and at the end of our incubation experiments.

Very strong but exchangeable organic ligand of cobalt in the marginal sea

AbstractCobalt (Co) is an important micronutrient for phytoplankton and its chemical speciation in the ocean is dominated by organic complexation. However, determination of the chemical speciation parameters of dissolved Co (dCo) in seawater is difficult because the ligands have conditional stability constants (K′) higher than those detectable by current forward titration methods. Furthermore, forward titration is not ideal when the ligand concentration is less than the dCo concentration in natural seawater, which is a probable scenario because excess ligands will bind more strongly to nickel instead. In this study, we successfully determined the concentrations and K′ of the dCo‐binding organic ligands in East China Sea using the reverse titration method. Our results indicate that organic complexes of dCo in the East China Sea are very strong but are kinetically exchangeable at high detection windows. Log K′ values of the ligands were between 16.6 and 17.1 in the Kuroshio waters and ranged from 19.7 to 20.9 in coastal waters. These ligands might be phytoplankton exudates or degradation products of cobalamin. Our findings suggest that an inert dCo fraction may be significant in coastal regions with high primary productivity. These inert dCo complexes were not detected within our detection window, which may lead to overestimation of the titration results.

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.

Phytoplankton exudates provide full nutrition to a subset of accompanying heterotrophic bacteria via carbon, nitrogen and phosphorus allocation.

Marine bacteria rely on phytoplankton exudates as carbon sources (DOCp). Yet, it is unclear to what extent phytoplankton exudates also provide nutrients such as phytoplankton-derived N and P (DONp, DOPp). We address these questions by mesocosm exudate addition experiments with spent media from the ubiquitous pico-cyanobacterium Prochlorococcus to bacterial communities in contrasting ecosystems in the Eastern Mediterranean - a coastal and an open-ocean, oligotrophic station with and without on-top additions of inorganic nutrients. Inorganic nutrient addition did not lower the incorporation of exudate DONp, nor did it reduce alkaline phosphatase activity, suggesting that bacterial communities are able to exclusively cover their nitrogen and phosphorus demands with organic forms provided by phytoplankton exudates. Approximately half of the cells in each ecosystem took up detectable amounts of Prochlorococcus-derived C and N, yet based on 16S rRNA sequencing different bacterial genera were responsible for the observed exudate utilization patterns. In the coastal community, several phylotypes of Aureimarina, Psychrosphaera and Glaciecola responded positively to the addition of phytoplankton exudates, whereas phylotypes of Pseudoalteromonas increased and dominated the open-ocean communities. Together, our results strongly indicate that phytoplankton exudates provide coastal and open-ocean bacterial communities with organic carbon, nitrogen and phosphorus, and that phytoplankton exudate serve a full-fledged meal for the accompanying bacterial community in the nutrient-poor eastern Mediterranean.

Phytoplankton exudates and lysates support distinct microbial consortia with specialized metabolic and ecophysiological traits

Blooms of marine phytoplankton fix complex pools of dissolved organic matter (DOM) that are thought to be partitioned among hundreds of heterotrophic microbes at the base of the food web. While the relationship between microbial consumers and phytoplankton DOM is a key component of marine carbon cycling, microbial loop metabolism is largely understood from model organisms and substrates. Here, we took an untargeted approach to measure and analyze partitioning of four distinct phytoplankton-derived DOM pools among heterotrophic populations in a natural microbial community using a combination of ecogenomics, stable isotope probing (SIP), and proteomics. Each 13C-labeled exudate or lysate from a diatom or a picocyanobacterium was preferentially assimilated by different heterotrophic taxa with specialized metabolic and physiological adaptations. Bacteroidetes populations, with their unique high-molecular-weight transporters, were superior competitors for DOM derived from diatom cell lysis, rapidly increasing growth rates and ribosomal protein expression to produce new relatively high C:N biomass. Proteobacteria responses varied, with relatively low levels of assimilation by Gammaproteobacteria populations, while copiotrophic Alphaproteobacteria such as the Roseobacter clade, with their diverse array of ABC- and TRAP-type transporters to scavenge monomers and nitrogen-rich metabolites, accounted for nearly all cyanobacteria exudate assimilation and produced new relatively low C:N biomass. Carbon assimilation rates calculated from SIP data show that exudate and lysate from two common marine phytoplankton are being used by taxonomically distinct sets of heterotrophic populations with unique metabolic adaptations, providing a deeper mechanistic understanding of consumer succession and carbon use during marine bloom events.

Resource Colimitation Drives Competition Between Phytoplankton and Bacteria in the Southern Ocean.

Across the Southern Ocean, phytoplankton growth is governed by iron and light, while bacterial growth is regulated by iron and labile dissolved organic carbon (LDOC). We use a mechanistic model to examine how competition for iron between phytoplankton and bacteria responds to changes in iron, light, and LDOC. Consistent with experimental evidence, increasing iron and light encourages phytoplankton dominance, while increasing LDOC and decreasing light favors bacterial dominance. Under elevated LDOC, bacteria can outcompete phytoplankton for iron, most easily under lower iron. Simulations reveal that bacteria are major iron consumers and suggest that luxury storage plays a key role in competitive iron uptake. Under seasonal conditions typical of the Southern Ocean, sources of LDOC besides phytoplankton exudation modulate the strength of competitive interactions. Continued investigations on the competitive fitness of bacteria in driving changes in primary production in iron‐limited systems will be invaluable in refining these results.

Enhanced formation of transparent exopolymer particles (TEP) under turbulence during phytoplankton growth

AbstractSmall-scale turbulence in the surface ocean is ubiquitous, influencing phytoplankton dynamics with consequences for energy flow. The underlying mechanisms that drive changes in phytoplankton dynamics under turbulence are not well constrained. We investigated growth of four phytoplankton species at different turbulence levels in oscillating grid tanks. We also measured transparent exopolymer particles (TEP) from phytoplankton exudates, which play a major role in biogeochemical fluxes in the ocean. Turbulence levels in the tanks reflected in situ conditions in surface waters from the open ocean to higher turbulent environments such as estuaries. Growth rates were unaffected by turbulence while TEP concentrations as xanthan gum (XG) equivalents normalized to algal cells showed generally higher levels in the high turbulence compared to the low turbulence treatments particularly during initial algal growth. Results from a mixing experiment without algal cells and XG also revealed enhanced formation of TEP-like particles under high mixing conditions, indicating that TEP formation in the phytoplankton turbulence treatments was mainly driven by physical processes, such as enhanced encounter rates of TEP-precursors under high mixing. Our results underline the importance of small-scale turbulence on TEP formation with possible consequences for particle aggregation and vertical carbon fluxes in the ocean.

Assessment of bacterial dependence on marine primary production along a northern latitudinal gradient.

Recent observations in polar marine waters have shown that a large fraction of primary production may be lost to respiration by planktonic bacteria due to very low bacterial growth efficiencies in cold waters. Here we report that sea temperature may be a key factor (but not the only one) influencing the interaction between bacteria and primary production in North Atlantic and Arctic waters, suggesting that low primary production rates could not sustain bacterial carbon demand in the coldest Arctic waters. The use of freshly produced phytoplankton exudate by bacteria in early- and mid-summer was assessed, together with the bacterial uptake of dissolved inorganic nitrogen (DIN=nitrate and ammonium), in surface waters along a latitudinal gradient from the North Sea to the Arctic sea ice. Bacterial production was independent of the low primary production measured in the coldest waters. Under these conditions, heterotrophic bacteria can consume a large fraction of DIN and N-rich organic matter, making them strong contributors to N fluxes in these waters.

Variability in the production of organic ligands, by Synechococcus PCC 7002, under different iron scenarios

Several Fe-uptake mechanisms suggest the importance of the presence of certain organic ligands in phytoplankton exudates. Here, it has been studied how Synechococcus (strain PCC 7002) acclimates to Fe-bioavailability, comparing the growth and organic exudation under two different Fe regimes. These cyanobacteria were incubated in UV-treated seawater supplemented only with major nutrients and two different iron scenarios (Low-Fe and High-Fe), without chelating agents, in order to analyze the organic ligands production. The levels of dissolved organic carbon (DOC) and two natural ligands (hydroxamic and phenolic moieties) were monitored. The responses in the organic extracellular release rates (ER), normalized per cell, were statistically analyzed considering Fe scenarios and different development stages. Growth of Synechococcus was significantly slower under Low-Fe treatment, suggesting that these cultures were iron limited compared to those flourished with higher levels of iron in the medium. Although the concentration of DOC increased to 127.13 ± 8.38 and 150.51 ± 8.59 μmol C L−1 under Low-Fe and High-Fe conditions, respectively, no-significant variations were found in the DOCER, among growth phases and iron bioavailability scenarios. Under High-Fe conditions, the production of hydroxamic ligands was inhibited, while the extracellular release rate of phenolic compounds decreased, regarding to Low-Fe conditions. Growth phases of Synechococcus also modified the extracellular release rates both of hydroxamic and phenolic moieties. The present study, therefore, demonstrates that iron availability and growth stages might be key parameters in regulating the release performance of extracellular Fe-specific organic ligands by cyanobacteria.

Ocean Acidification Experiments in Large-Scale Mesocosms Reveal Similar Dynamics of Dissolved Organic Matter Production and Biotransformation

Dissolved organic matter (DOM) represents a major reservoir of carbon in the oceans. Environmental stressors such as ocean acidification (OA) potentially affect DOM production and degradation processes, e.g. phytoplankton exudation or microbial uptake and biotransformation of molecules. Resulting changes in carbon storage capacity of the ocean, thus, may cause feedbacks on the global carbon cycle. Previous experiments studying OA effects on the DOM pool under natural conditions, however, were mostly conducted in temperate and coastal eutrophic areas. Here, we report on OA effects on the existing and newly produced DOM pool during an experiment in the subtropical North Atlantic Ocean at the Canary Islands during an (1) oligotrophic phase and (2) after simulated deep water upwelling. The last is a frequently occurring event in this region controlling nutrient and phytoplankton dynamics. We manipulated nine large-scale mesocosms with a gradient of pCO2 ranging from ~350 up to ~1030 µatm and monitored the DOM molecular composition using ultrahigh-resolution mass spectrometry via Fourier-transform ion cyclotron resonance mass spectrometry (FT-ICR-MS). An increase of 37 µmol L-1 DOC was observed in all mesocosms during a phytoplankton bloom induced by simulated upwelling. Indications for enhanced DOC accumulation under elevated CO2 became apparent during a phase of nutrient recycling towards the end of the experiment. The production of DOM was reflected in changes of the molecular DOM composition. Out of the 7,212 molecular formulae, which were detected throughout the experiment, ~50% correlated significantly in mass spectrometric signal intensity with cumulative bacterial protein production and are likely a product of microbial transformation. However, no differences in the produced compounds were found with respect to CO2 levels. Comparing the results of this experiment with a comparable OA experiment in the Swedish Gullmar Fjord, reveals similar succession patterns for individual compound pools during a phytoplankton bloom and subsequent accumulation of these compounds were observed. The similar behavior of DOM production and biotransformation during and following a phytoplankton bloom irrespective of plankton community composition and CO2 treatment provides novel insights into general dynamics of the marine DOM pool.

Elevated pCO2 enhances bacterioplankton removal of organic carbon.

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.

Combined Carbohydrates Support Rich Communities of Particle-Associated Marine Bacterioplankton.

Carbohydrates represent an important fraction of labile and semi-labile marine organic matter that is mainly comprised of exopolymeric substances derived from phytoplankton exudation and decay. This study investigates the composition of total combined carbohydrates (tCCHO; >1 kDa) and the community development of free-living (0.2–3 μm) and particle-associated (PA) (3–10 μm) bacterioplankton during a spring phytoplankton bloom in the southern North Sea. Furthermore, rates were determined for the extracellular enzymatic hydrolysis that catalyzes the initial step in bacterial organic matter remineralization. Concentrations of tCCHO greatly increased during bloom development, while the composition showed only minor changes over time. The combined concentration of glucose, galactose, fucose, rhamnose, galactosamine, glucosamine, and glucuronic acid in tCCHO was a significant factor shaping the community composition of the PA bacteria. The richness of PA bacteria greatly increased in the post-bloom phase. At the same time, the increase in extracellular β-glucosidase activity was sufficient to explain the observed decrease in tCCHO, indicating the efficient utilization of carbohydrates by the bacterioplankton community during the post-bloom phase. Our results suggest that carbohydrate concentration and composition are important factors in the multifactorial environmental control of bacterioplankton succession and the enzymatic hydrolysis of organic matter during phytoplankton blooms.

Biological or microbial carbon pump? The role of phytoplankton stoichiometry in ocean carbon sequestration

Once fixed by photosynthesis carbon becomes part of the marine food web. The fate of this carbon has two possible outcomes, it may be respired and released back to the ocean and potentially to the atmosphere as CO2 or retained in the ocean interior and/or marine sediments for extended time scales. The most important biologically mediated processes responsible for long-term carbon storage in the ocean are the biological carbon pump (BCP) and the microbial carbon pump (MCP). While acting simultaneously in the ocean, the balance between these two mechanisms is thought to vary depending on the trophic state of the environment. Using previously published formulations, we propose a modelling framework to simulate variability in the MCP:BCP ratio as a function of external nutrients. Our results suggest that the role of the MCP might become more significant under future climate change conditions where increased stratification enhances the oligotrophic nature of the surface ocean. Based on these model results, we propose a conceptual framework in which the internal stoichiometry of phytoplankton, modulating both grazing pressure and dissolved organic matter production (via phytoplankton exudation), plays a crucial role in regulating the MCP:BCP ratio.

Spatio-Temporal Interdependence of Bacteria and Phytoplankton during a Baltic Sea Spring Bloom.

In temperate systems, phytoplankton spring blooms deplete inorganic nutrients and are major sources of organic matter for the microbial loop. In response to phytoplankton exudates and environmental factors, heterotrophic microbial communities are highly dynamic and change their abundance and composition both on spatial and temporal scales. Yet, most of our understanding about these processes comes from laboratory model organism studies, mesocosm experiments or single temporal transects. Spatial-temporal studies examining interactions of phytoplankton blooms and bacterioplankton community composition and function, though being highly informative, are scarce. In this study, pelagic microbial community dynamics (bacteria and phytoplankton) and environmental variables were monitored during a spring bloom across the Baltic Proper (two cruises between North Germany to Gulf of Finland). To test to what extent bacterioplankton community composition relates to the spring bloom, we used next generation amplicon sequencing of the 16S rRNA gene, phytoplankton diversity analysis based on microscopy counts and population genotyping of the dominating diatom Skeletonema marinoi. Several phytoplankton bloom related and environmental variables were identified to influence bacterial community composition. Members of Bacteroidetes and Alphaproteobacteria dominated the bacterial community composition but the bacterial groups showed no apparent correlation with direct bloom related variables. The less abundant bacterial phyla Actinobacteria, Planctomycetes, and Verrucomicrobia, on the other hand, were strongly associated with phytoplankton biomass, diatom:dinoflagellate ratio, and colored dissolved organic matter (cDOM). Many bacterial operational taxonomic units (OTUs) showed high niche specificities. For example, particular Bacteroidetes OTUs were associated with two distinct genetic clusters of S. marinoi. Our study revealed the complexity of interactions of bacterial taxa with inter- and intraspecific genetic variation in phytoplankton. Overall, our findings imply that biotic and abiotic factors during spring bloom influence bacterial community dynamics in a hierarchical manner.

Heterotrophic Bacteria Show Weak Competition for Nitrogen in Mediterranean Coastal Waters (Thau Lagoon) in Autumn.

The importance of heterotrophic bacteria relative to phytoplankton in the uptake of ammonium and nitrate was studied in Mediterranean coastal waters (Thau Lagoon) during autumn, when the Mediterranean Sea received the greatest allochthonous nutrient loads. Specific inhibitors and size-fractionation methods were used in combination with isotopic (15)N tracers. NO3 (-) and NH4 (+) uptake was dominated by phytoplankton (60 % on average) during the study period, which included a flood event. Despite lower biomass specific NH4 (+) and NO3 uptake rates, free-living heterotrophic bacteria contributed significantly (>30 %) to total microbial NH4 (+) and NO3 (-) uptake rates in low chlorophyll waters. Under these conditions, heterotrophic bacteria may be responsible for more than 50 % of primary production, using very little freshly produced phytoplankton exudates. In low chlorophyll coastal waters as reported during the present 3-month study, the heterotrophic bacteria seemed to depend to a greater extent on allochthonous N and C substrates than on autochthonous substrates derived from phytoplankton.

Differentiating humic and algal surface active substances in coastal waters by their pH-dependent adsorption behaviour

Sinusoidal alternating current (ac) voltammetry is a long-established and very sensitive technique for the determination of surface active substances (SAS) in natural waters. However, different classes of SAS cannot be distinguished from the shape of their ac current vs. potential (i–E) curves alone. Here we describe an effective method for quantifying the mixed pools of marine algal exudates and aquatic humic substances present in coastal waters. The method involves monitoring the adsorption kinetics of naturally occurring SAS at two different pH values, here chosen to be pH3.5 and 7.0. We found that a simple rectangular hyperbola to describe the surface coverage of the mercury drop versus time yielded an excellent fit from which a kinetic adsorption constant k (s−1) could be derived. In addition, k increased linearly with concentration of the adsorbing species. Calibration plots of k vs. concentration for two model substances, i.e. Nordic Reservoir NOM and diatom culture exudates, were obtained. The value of k was found to be a sensitive function of pH between pH3.5 and 7.0 for humic substances, but not for phytoplankton exudates. This provided the basis for resolving the concentrations of ‘humic-like’ and ‘algal-like’ SAS in natural samples collected along a transect extending from the mouth of a humic-rich river (S=5) to the far field region of the river plume (S>34). Concentration–salinity relationships indicated a terrestrial source of humic SAS and a marine source of algal SAS, as expected. Calculated dissolved organic carbon ([DOC]) concentrations were in good agreement with [DOC] measured directly.

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.

Bacterial carbon dependence on freshly produced phytoplankton exudates under different nutrient availability and grazing pressure conditions in coastal marine waters.

The effects of grazing pressure and inorganic nutrient availability on the direct carbon transfer from freshly produced phytoplankton exudates to heterotrophic bacteria biomass production were studied in Mediterranean coastal waters. The short-term incorporation of ¹³C (H¹³CO₃) in phytoplankton and bacterial lipid biomarkers was measured as well as the total bacterial carbon production (BP), viral lysis and the microbial community structure under three experimental conditions: (1) High inorganic Nutrient and High Grazing (HN + HG), (2) High inorganic Nutrient and Low Grazing (HN + LG) and (3) under natural in situ conditions with Low inorganic Nutrient and High Grazing (LN + HG) during spring. Under phytoplankton bloom conditions (HN + LG), the bacterial use of freshly produced phytoplankton exudates as a source of carbon, estimated from ¹³C enrichment of bacterial lipids, contributed more than half of the total bacterial production. However, under conditions of high grazing pressure on phytoplankton with or without the addition of inorganic nutrients (HN + HG and LN + HG), the ¹³C enrichment of bacterial lipids was low compared with the high total bacterial production. BP therefore seems to depend mainly on freshly produced phytoplankton exudates during the early phase of phytoplankton bloom period. However, BP seems mainly relying on recycled carbon from viral lysis and predators under high grazing pressure.