scholarly journals Strong stimulation of N<sub>2</sub> fixation in oligotrophic Mediterranean Sea: results from dust addition in large in situ mesocosms

2013 ◽  
Vol 10 (11) ◽  
pp. 7333-7346 ◽  
Author(s):  
C. Ridame ◽  
C. Guieu ◽  
S. L'Helguen

Abstract. The response of N2 (dinitrogen) fixation to contrasted (wet and dry) Saharan dust deposition was studied in the framework of the DUNE project (a DUst experiment in a low-Nutrient, low-chlorophyll Ecosystem) during which realistic simulations of dust deposition (10 g m−2) into large mesocosms (52 m3) were performed. Three distinct experimental dust additions were conducted in June 2008 (DUNE-1-P: simulation of a wet deposition, DUNE-1-Q: simulation of a dry deposition) and 2010 (DUNE-2-R: simulation of 2 successive wet depositions) in the northwestern oligotrophic Mediterranean Sea. Here we show that wet and dry dust deposition induced a rapid (24 h or 48 h after dust additions), strong (from 2- to 5.3-fold) and long (at least 4–6 days duration) increase in N2 fixation, indicating that both wet and dry Saharan dust deposition was able to relieve efficiently the nutrient limitation(s) of N2 fixation. This means in particular that N2 fixation activity was not inhibited by the significant input of nitrate associated with the simulated wet deposition (~ 9 mmol NO3− m−2). The input of new nitrogen associated with N2 fixation was negligible relative to the atmospheric NO3− input associated with the dust. The contribution of N2 fixation to primary production was negligible (≤ 1%) before and after dust addition in all experiments, indicating that N2 fixation was a poor contributor to the nitrogen demand for primary production. Despite the stimulation of N2 fixation by dust addition, the rates remained low, and did not significantly change the contribution of N2 fixation to new production since only a maximum contribution of 10% was observed. The response of N2 fixation by diazotrophs and CO2 fixation by the whole phytoplankton community suggests that these metabolic processes were limited or co-limited by different nutrients. With this novel approach, which allows us to study processes as a function of time while atmospheric particles are sinking, we show that new atmospheric nutrients associated with Saharan dust pulses do significantly stimulate N2 fixation in the Mediterranean Sea and that N2 fixation is not a key process in the carbon cycle in such oligotrophic environments.

2014 ◽  
Vol 11 (1) ◽  
pp. 753-796 ◽  
Author(s):  
C. Ridame ◽  
J. Dekaezemacker ◽  
C. Guieu ◽  
S. Bonnet ◽  
S. L'Helguen ◽  
...  

Abstract. The response of the phytoplanktonic community (primary production and algal biomass) to contrasted Saharan dust events (wet and dry deposition) was studied in the framework of the DUNE "a DUst experiment in a low-Nutrient, low-chlorophyll Ecosystem" project. We simulated realistic dust deposition events (10 g m−2) into large mesocosms (52 m3). Three distinct experimental dust additions were conducted in June 2008 (DUNE-1-P: simulation of a wet deposition, DUNE-1-Q: simulation of a dry deposition) and 2010 (DUNE-2-R1, -R2: simulation of 2 successive wet depositions) in the northwestern oligotrophic Mediterranean Sea. No changes in primary production (PP) and chlorophyll a concentration (Chl a) were observed after a dry deposition event while a wet deposition event resulted in a rapid (24 h after dust additions), strong (up 2.4 fold) and long (at least a week duration) increase in PP and Chl a. We show that in addition to being a source of dissolved inorganic phosphorus (DIP), simulated wet deposition events were also a significant source of NO3− (net increases up to +9.8 μM NO3− at 0.1 m depth) to the nutrient depleted surface waters due to cloud processes and mixing with anthropogenic species such as HNO3. The dry deposition event was shown to be a negligible source of NO3−. By transiently increasing DIP and NO3− concentrations in P-N starved surface waters, wet deposition of Saharan dust was able to relieve the potential N or NP co-limitation of the phytoplanktonic activity. Due to the higher input of NO3− relative to DIP, a wet deposition event resulted in a strong increase in the NO3−/DIP ratio from initially < 6 to over 150 at the end of the DUNE-2-R1 experiment suggesting a switch from an initial N or NP co-limitation towards a severe P limitation. We also show that the contribution of new production to PP increased after wet dust deposition events from initially 15% to 60–70% 24 h after seeding, indicating a switch from a regenerated-production based system to a new-production based system. DUNE experiments show that wet and dry dust deposition events induce contrasted responses of the phytoplanktonic community due to differences in the atmospheric supply of bioavailable new nutrients. Our results from original mesocosm experiments demonstrate that atmospheric dust wet deposition greatly influences primary productivity and algal biomass in LNLC environments, changes nutrient stocks and alters the NO3−/DIP ratio leading to a switch in the nutrient limitation of the phytoplanktonic activity.


2014 ◽  
Vol 11 (17) ◽  
pp. 4783-4800 ◽  
Author(s):  
C. Ridame ◽  
J. Dekaezemacker ◽  
C. Guieu ◽  
S. Bonnet ◽  
S. L'Helguen ◽  
...  

Abstract. The response of the phytoplanktonic community (primary production and algal biomass) to contrasted Saharan dust events (wet and dry deposition) was studied in the framework of the DUNE ("a DUst experiment in a low-Nutrient, low-chlorophyll Ecosystem") project. We simulated realistic dust deposition events (10 g m−2) into large mesocosms (52 m3). Three distinct dust addition experiments were conducted in June 2008 (DUNE-1-P: simulation of a wet deposition; DUNE-1-Q: simulation of a dry deposition) and 2010 (DUNE-2-R1 and DUNE-2-R2: simulation of two successive wet depositions) in the northwestern oligotrophic Mediterranean Sea. No changes in primary production (PP) and chlorophyll a concentrations (Chl a) were observed after a dry deposition event, while a wet deposition event resulted in a rapid (24 h after dust addition), strong (up to 2.4-fold) and long (at least a week in duration) increase in PP and Chl a. We show that, in addition to being a source of dissolved inorganic phosphorus (DIP), simulated wet deposition events were also a significant source of nitrate (NO3−) (net increases up to +9.8 μM NO3− at 0.1 m in depth) to the nutrient-depleted surface waters, due to cloud processes and mixing with anthropogenic species such as HNO3. The dry deposition event was shown to be a negligible source of NO3−. By transiently increasing DIP and NO3- concentrations in N–P starved surface waters, wet deposition of Saharan dust was able to relieve the potential N or NP co-limitation of the phytoplanktonic activity. Due to the higher input of NO3− relative to DIP, and taking into account the stimulation of the biological activity, a wet deposition event resulted in a strong increase in the NO3−/DIP ratio, from initially less than 6, to over 150 at the end of the DUNE-2-R1 experiment, suggesting a switch from an initial N or NP co-limitation towards a severe P limitation. We also show that the contribution of new production to PP strongly increased after wet dust deposition events, from initially 15% to 60–70% 24 h after seeding, indicating a switch from a regenerated-production based system to a new-production based system. DUNE experiments show that wet and dry dust deposition events induce contrasting responses of the phytoplanktonic community due to differences in the atmospheric supply of bioavailable new nutrients. Our results from original mesocosm experiments demonstrate that atmospheric dust wet deposition greatly influences primary productivity and algal biomass in LNLC environments through changes in the nutrient stocks, and alters the NO3−/DIP ratio, leading to a switch in the nutrient limitation of the phytoplanktonic activity.


2013 ◽  
Vol 10 (6) ◽  
pp. 10581-10613 ◽  
Author(s):  
C. Ridame ◽  
C. Guieu ◽  
S. L'Helguen

Abstract. The response of N2 fixation to contrasted (wet and dry) Saharan dust deposition was studied in the framework of the DUNE project "a DUst experiment in a low-Nutrient, low-chlorophyll Ecosystem" during which realistic simulations of dust deposition (10 g m


2014 ◽  
Vol 11 (19) ◽  
pp. 5621-5635 ◽  
Author(s):  
C. Guieu ◽  
C. Ridame ◽  
E. Pulido-Villena ◽  
M. Bressac ◽  
K. Desboeufs ◽  
...  

Abstract. By bringing new nutrients and particles to the surface ocean, atmospheric deposition impacts biogeochemical cycles. The extent to which those changes are modifying the carbon balance in oligotrophic environments such as the Mediterranean Sea that receives important Saharan dust fluxes is unknown. The DUNE (DUst experiment in a low Nutrient, low chlorophyll Ecosystem) project provides the first attempt to evaluate the changes induced in the carbon budget of a large body of oligotrophic waters after simulated Saharan dust wet or dry deposition events, allowing us to measure (1) the metabolic fluxes while the particles are sinking and (2) the particulate organic carbon export. Here we report the results for the three distinct artificial dust seeding experiments simulating wet or dry atmospheric deposition onto large mesocosms (52 m3) that were conducted in the oligotrophic waters of the Mediterranean Sea in the summers of 2008 and 2010. Although heterotrophic bacteria were found to be the key players in the response to dust deposition, net primary production increased about twice in case of simulated wet deposition (that includes anthropogenic nitrogen). The dust deposition did not produce a shift in the metabolic balance as the tested waters remained net heterotrophic (i.e., net primary production to bacteria respiration ratio <1) and in some cases the net heterotrophy was even enhanced by the dust deposition. The change induced by the dust addition on the total organic carbon pool inside the mesocosm over the 7 days of the experiments, was a carbon loss dominated by bacteria respiration that was at least 5–10 times higher than any other term involved in the budget. This loss of organic carbon from the system in all the experiments was particularly marked after the simulation of wet deposition. Changes in biomass were mostly due to an increase in phytoplankton biomass but when considering the whole particulate organic carbon pool it was dominated by the organic carbon aggregated to the lithogenic particles still in suspension in the mesocosm at the end of the experiment. Assuming that the budget is balanced, the dissolved organic carbon (DOC) pool was estimated by the difference between the total organic carbon and the particulate organic carbon (POC) pool. The partitioning between dissolved and particulate organic carbon was dominated by the dissolved pool with a DOC consumption over 7 days of ∼1 μmol C L−1 d−1 (dry deposition) to ∼2–5 μmol C L−1 d−1 (wet deposition). This consumption in the absence of any allochthonous inputs in the closed mesocosms meant a small <10% decrease of the initial DOC stock after a dry deposition but a ∼30–40% decrease of the initial DOC stock after wet deposition. After wet deposition, the tested waters, although dominated by heterotrophy, were still maintaining a net export (corrected from controls) of particulate organic carbon (0.5 g in 7 days) even in the absence of allochthonous carbon inputs. This tentative assessment of the changes in carbon budget induced by a strong dust deposition indicates that wet deposition by bringing new nutrients has higher impact than dry deposition in oligotrophic environments. In the western Mediterranean Sea, the mineral dust deposition is dominated by wet deposition and one perspective of this work is to extrapolate our numbers to time series of deposition during similar oligotrophic conditions to evaluate the overall impact on the carbon budget at the event and seasonal scale in the surface waters of the northwestern Mediterranean Sea. These estimated carbon budgets are also highlighting the key processes (i.e., bacterial respiration) that need to be considered for an integration of atmospheric deposition in marine biogeochemical modeling.


2011 ◽  
Vol 8 (2) ◽  
pp. 2629-2657 ◽  
Author(s):  
C. Ridame ◽  
M. Le Moal ◽  
C. Guieu ◽  
E. Ternon ◽  
I. C. Biegala ◽  
...  

Abstract. A better understanding of the factors controlling N2 fixation is a pre-requisite for improving our knowledge on the contribution of N2 fixation in the nitrogen cycling in the Mediterranean Sea. Trace-metal clean nutrient/dust additions bioassays (+P, +PFe, +dust) were performed at three stations located in the western, central and eastern Mediterranean Sea, in summer 2008 as part of the BOUM cruise. The main goals were to investigate the nutrient factor(s) limiting N2 fixation (uptake of 15N2) and to evaluate the potential impact of a Saharan dust event on this biological process during the stratification period. Initially, surface waters at the three stations were DIP-depleted (<10 nM) while the DFe concentrations were relatively high (from 1.2 to 2.3 nM) most likely due to atmospheric iron accumulation in the surface mixed layer. At all stations, Saharan dust input relieved the ambient nutrient limitation of diazotrophic activity as demonstrated by the strong stimulation of N2 fixation (from x2.3 to x5.3). The highest dust stimulation of N2 fixation was recorded at the station located in the eastern basin (x5.3). The responses of diazotrophic activity to nutrients addition were contrasted at the sampled stations suggesting a spatial variability of the factor controlling N2 fixation over the whole basin. At all stations, N2 fixation was not limited by Fe nor co-limited by P and Fe. At the western station, N2 fixation was DIP limited while at the eastern one, N2 fixation was first DIP limited then was limited by one or several chemical element(s) released by dust. Our results demonstrated that a Saharan dust input was able to relieve the successive on-going N2 fixation limitations. Very interestingly, at the station located in the central basin, N2 fixation was not limited by the availability of P yet it was strongly stimulated by dust additions (up to x3.1). A chemical element or a combination of several, released by the added dust may have been responsible for the observed stimulations of N2 fixation. These results indicated that Saharan dust pulses to the surface Mediterranean waters, in addition to P and Fe, could be a source of chemical(s) element(s) that are necessary for metabolic processes and therefore influence rates of N2 fixation.


2021 ◽  
Author(s):  
Céline Ridame ◽  
Julie Dinasquet ◽  
Søren Hallstrøm ◽  
Estelle Bigeard ◽  
Lasse Riemann ◽  
...  

Abstract. N2 fixation rates were measured in the 0–1000 m layer at 13 stations located in the open western and central Mediterranean Sea (MS) during the PEACETIME cruise (late spring 2017). While the spatial variability of N2 fixation was not related to Fe, P nor N stocks, the surface composition of the diazotrophic community indicated a strong eastward increasing longitudinal gradient for the relative abundance of non-cyanobacterial diazotrophs (NCD) (mainly γ-Proteobacteria) and conversely eastward decreasing for UCYN-A (mainly -A1 and -A3) as did N2 fixation rates. UCYN-A4 and A3 were identified for the first time in the MS. The westernmost station influenced by Atlantic waters, and characterized by highest stocks of N and P, displayed a patchy distribution of diazotrophic activity with an exceptionally high rate in the euphotic layer of 72.1 nmol N L−1 d−1, which could support up to 19 % of primary production. At this station at 1 %PAR depth, UCYN-A4 represented up to 94 % of the diazotrophic community. These in situ observations of higher UCYN-A relative abundance in nutrient rich stations while NCD increased in the more oligotrophic stations, suggest that the nutrient conditions could determine the composition of the diazotrophic communities and in turn the N2 fixation rates. The impact of Saharan dust deposition on N2 fixation and diazotrophic communities was also investigated, under present and future projected conditions of temperature and pH during short term (3–4 days) experiments at three stations. New nutrients from simulated dust deposition triggered a significant stimulation of N2 fixation (from 41 % to 565 %). The strongest increase in N2 fixation was observed at the stations dominated by NCD and did not lead on this short time scale to change in the diazotrophic community composition. Under projected future conditions, N2 fixation was either exacerbated or unchanged, in that later case this was probably due to a too low nutrient bioavailability or an increased grazing pressure. The future warming and acidification likely benefited NCD (Pseudomonas) and UCYN-A2 while disadvantaged UCYN-A3 without knowing which effect (alone or in combination) is the driver, especially since we do not know the temperature optima of these species not yet cultivated as well as the effect of acidification.


2021 ◽  
Author(s):  
Karine Desboeufs ◽  
Franck Fu ◽  
Matthieu Bressac ◽  
Antonio Tovar-Sánchez ◽  
Sylvain Triquet ◽  
...  

Abstract. This study reports the only recent characterisation of two contrasted wet deposition events collected during the PEACETIME cruise in the Mediterranean open seawater, and their impact on trace metals (TMS) marine stocks. Rain samples were analysed for Al, 12 trace metals (TMs hereafter, including Co, Cd, Cr, Cu, Fe, Mn, Mo, Ni, Pb, Ti, V and Zn) and nutrients (N, P, DOC) concentrations. The first rain sample collected in the Ionian Sea (rain ION) was a wet typical regional background deposition event whereas the second rain collected in the Algerian Basin (rain FAST) was a Saharan dust wet deposition. The concentrations of TMs in the two rain samples were significantly lower compared to concentrations in rains collected at coastal sites reported in the literature, suggesting either less anthropogenic influence in the remote Mediterranean environment, or decreased emissions during the last decades in the Mediterranean Sea. The TMs inventories in the surface microlayer and mixed layer (0–20 m) at ION and FAST stations before and after the events, compared to atmospheric fluxes, showed that the atmospheric inputs were a significant source of particulate TMs for both layers. At the scale of the western and central Mediterranean, the atmospheric inputs were of the same order of magnitude as marine stocks within the ML for dissolved Fe, Co and Zn, underlining the role of the atmosphere in their biogeochemical cycle in the stratified Mediterranean Sea. In case of intense wet dust deposition event, the contribution of atmospheric inputs could be critical for dissolved stocks of the majority of TMs.


2014 ◽  
Vol 11 (1) ◽  
pp. 1707-1738 ◽  
Author(s):  
C. Guieu ◽  
C. Ridame ◽  
E. Pulido-Villena ◽  
M. Bressac ◽  
K. Desboeufs ◽  
...  

Abstract. By bringing new nutrients and particles to the surface ocean, atmospheric deposition impacts biogeochemical cycles. The extent to which those changes are modifying the carbon balance in oligotrophic environments such as the Mediterranean Sea that receives important Saharan dust fluxes is unknown. DUNE project provides the first attempt to evaluate the changes induced in the carbon budget of an oligotrophic system after simulated Saharan dust wet and dry deposition events. Here we report the results for the 3 distinct artificial dust seeding experiments in large mesocosms that were conducted in the oligotrophic waters of the Mediterranean Sea in summer 2008 and 2010. Simultaneous measurements of the metabolic rates (C fixation, C respiration) in the water column have shown that the dust deposition did not change drastically the metabolic balance as the tested waters remained net heterotroph (i.e. net primary production to bacteria respiration ratio < 1) and in some cases the net heterotrophy was even enhanced by the dust deposition. Considering the different terms of the carbon budget, we estimate that it was balanced with a dissolved organic carbon (DOC) consumption of at least 10% of the initial stock. This corresponds to a fraction of the DOC stock of the surface mixed layer that consequently will not be exported during the winter mixing. Although heterotrophic bacteria were found to be the key players in the response to dust deposition, net primary production increased about twice in case of simulated wet deposition (that includes anthropogenic nitrogen) and a small fraction of particulate organic carbon was still exported. Our estimated carbon budgets are an important step forward in the way we understand dust deposition and associated impacts on the oceanic cycles. They are providing knowledge about the key processes (i.e. bacteria respiration, aggregation) that need to be considered for an integration of atmospheric deposition in marine biogeochemical modeling.


2011 ◽  
Vol 8 (9) ◽  
pp. 2773-2783 ◽  
Author(s):  
C. Ridame ◽  
M. Le Moal ◽  
C. Guieu ◽  
E. Ternon ◽  
I. C. Biegala ◽  
...  

Abstract. A better understanding of the factors controlling N2 fixation is a pre-requisite for improving our knowledge on the contribution of N2 fixation process in the nitrogen cycling. Trace-metal clean nutrient/dust addition bioassays (+P, +PFe, +dust) were performed at three stations located in the western, central and eastern Mediterranean Sea, in summer 2008 as part of the BOUM cruise. The main goals were (1) to investigate the nutrient factor(s) limiting N2 fixation (uptake of 15N2) and (2) to evaluate the potential impact of a Saharan dust event on this biological process during the stratification period. Initially, surface waters at the three stations were DIP-depleted (<10 nM) while the DFe concentrations were relatively high (from 1.2 to 2.3 nM) most likely due to atmospheric iron accumulation in the surface mixed layer. At all stations, Saharan dust input relieved the ambient nutrient limitation of the diazotrophic activity as demonstrated by the strong stimulation of N2 fixation (from 130 % to 430 %). The highest dust stimulation of N2 fixation was recorded at the station located in the eastern basin. The response of diazotrophic activity to nutrient additions was variable between the sampled stations suggesting a spatial variability of the factor controlling N2 fixation over the whole basin. At all stations, N2 fixation was not limited by Fe nor co-limited by P and Fe. At the western station, N2 fixation was DIP limited while at the eastern one, N2 fixation was first DIP limited, then was limited by one or several chemical element(s) released by dust. Our results demonstrated that a Saharan dust input was able to relieve these successive on going limitations. Very interestingly, at the station located in the central basin, N2 fixation was not limited by the availability of P yet it was strongly stimulated by dust addition (x3.1). A chemical element or a combination of several, released by the added dust may have been responsible for the observed stimulations of N2 fixation. These results indicated that Saharan dust pulses to the surface Mediterranean waters, in addition to P and Fe, could be a source of chemical(s) element(s) that are necessary for metabolic processes and therefore influence rates of N2 fixation.


2014 ◽  
Vol 11 (2) ◽  
pp. 425-442 ◽  
Author(s):  
C. Guieu ◽  
F. Dulac ◽  
C. Ridame ◽  
P. Pondaven

Abstract. The main goal of project DUNE was to estimate the impact of atmospheric deposition on an oligotrophic ecosystem based on mesocosm experiments simulating strong atmospheric inputs of eolian mineral dust. Our mesocosm experiments aimed at being representative of real atmospheric deposition events onto the surface of oligotrophic marine waters and were an original attempt to consider the vertical dimension after atmospheric deposition at the sea surface. This introductory paper describes the objectives of DUNE and the implementation plan of a series of mesocosm experiments conducted in the Mediterranean Sea in 2008 and 2010 during which either wet or dry and a succession of two wet deposition fluxes of 10 g m−2 of Saharan dust have been simulated based on the production of dust analogs from erodible soils of a source region. After the presentation of the main biogeochemical initial conditions of the site at the time of each experiment, a general overview of the papers published in this special issue is presented. From laboratory results on the solubility of trace elements in dust to biogeochemical results from the mesocosm experiments and associated modeling, these papers describe how the strong simulated dust deposition events impacted the marine biogeochemistry. Those multidisciplinary results are bringing new insights into the role of atmospheric deposition on oligotrophic ecosystems and its impact on the carbon budget. The dissolved trace metals with crustal origin – Mn, Al and Fe – showed different behaviors as a function of time after the seeding. The increase in dissolved Mn and Al concentrations was attributed to dissolution processes. The observed decrease in dissolved Fe was due to scavenging on sinking dust particles and aggregates. When a second dust seeding followed, a dissolution of Fe from the dust particles was then observed due to the excess Fe binding ligand concentrations present at that time. Calcium nitrate and sulfate were formed in the dust analog for wet deposition following evapocondensation with acids for simulating cloud processing by polluted air masses under anthropogenic influence. Using a number of particulate tracers that were followed in the water column and in the sediment traps, it was shown that the dust composition evolves after seeding by total dissolution of these salts. This provided a large source of new dissolved inorganic nitrogen (DIN) in the surface waters. In spite of this dissolution, the typical inter-elemental ratios in the particulate matter, such as Ti / Al or Ba / Al, are not affected during the dust settling, confirming their values as proxies of lithogenic fluxes or of productivity in sediment traps. DUNE experiments have clearly shown the potential for Saharan wet deposition to modify the in situ concentrations of dissolved elements of biogeochemical interest such as Fe and also P and N. Indeed, wet deposition yielded a transient increase in dissolved inorganic phosphorus (DIP) followed by a very rapid return to initial conditions or no return to initial conditions when a second dust seeding followed. By transiently increasing DIP and DIN concentrations in P- and N-starved surface waters of the Mediterranean Sea, wet deposition of Saharan dust can likely relieve the potential P and/or N limitation of biological activity; this has been directly quantified in terms of biological response. Wet deposition of dust strongly stimulated primary production and phytoplanktonic biomass during several days. Small phytoplankton (< 3 μm) was more stimulated after the first dust addition, whereas the larger size class (> 3 μm) significantly increased after the second one, indicating that larger-sized cells need further nutrient supply in order to be able to adjust their physiology and compete for resource acquisition and biomass increase. Among the microorganisms responding to the atmospheric inputs, diazotrophs were stimulated by both wet and dry atmospheric deposition, although N2 fixation was shown to be only responsible for a few percent of the induced new production. Dust deposition modified the bacterial community structure by selectively stimulating and inhibiting certain members of the bacterial community. The microbial food web dynamics were strongly impacted by dust deposition. The carbon budget indicates that the net heterotrophic character (i.e., ratio of net primary production to bacteria respiration < 1) of the tested waters remained (or was even increased) after simulated wet or dry deposition despite the significant stimulation of autotrophs after wet events. This indicates that the oligotrophic tested waters submitted to dust deposition are a net CO2 source. Nonetheless, the system was able to export organic material, half of it being associated with lithogenic particles through aggregation processes between lithogenic particles and organic matter. These observations support the "ballast" hypothesis and suggest that this "lithogenic carbon pump" could represent a major contribution of the global carbon export to deep waters in areas receiving high rates of atmospheric deposition. Furthermore, a theoretical microbial food web model showed that, all other things being equal, carbon, nitrogen and phosphorus stoichiometric mismatch along the food chain can have a substantial impact on the ecosystem response to nutrient inputs from dusts, with changes in the biomass of all biological compartments by a factor of ~ 2–4, and shifts from net autotrophy to net heterotrophy. Although the model was kept simple, it highlights the importance of stoichiometric constrains on the dynamics of microbial food webs.


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