Dust deposition in an oligotrophic marine environment: impact on the carbon budget

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 re-ceives important Saharan dust ﬂuxes is unknown. DUNE project provides the ﬁrst at- 5 tempt 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) 10 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 di ﬀ erent terms of the carbon budget, we estimate that it was balanced with a dissolved organic carbon (DOC) 15 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 20 of particulate organic carbon was still exported. Our estimated carbon budgets are


Introduction
Atmospheric deposition provides new nutrients to the marine environment and, in a number of oligotrophic environments submitted to significant deposition, those new nutrients are likely able to relieve on going (co)limitations by providing available nitrogen, phosphorus, iron and other micronutrients (de Leeuw et al., 2013). The response 5 of the biota to such input (natural, polluted or mixed aerosols) has been explored in a number of experimental artificial aerosols addition and in situ observations. In a recent paper, Guieu et al. (2014a) have synthetized the results from 26 of those studies showing that biota in Low-Nutrients and Low-Chlorophyll (LNLC) areas positively responds to aerosol addition, with bacterial production and N 2 fixation showing the 10 strongest responses. Increases in chlorophyll a (Chl a) have also been seen but to a lesser extent. In fact, this synthesis shows that changes in biological standing stocks tend to be smaller than changes in metabolic rates (i.e. chlorophyll a vs. primary production, and bacterial abundance vs. bacterial respiration and/or production). Indeed, this could be due to the effect of grazing by zooplankton on phytoplankton and bacteria, 15 (i.e. Bonnet et al., 2005;Herut et al., 2005;Marañón et al., 2010) which may increase turnover at the expense of stocks and rapidly propagate responses through the whole food web. This may result in an experimental artifact in that Chl a concentration may increase in experiments where grazers are filtered, but remains unchanged in natural waters, possibly explaining the variable and low response of satellite-derived chloro-20 phyll signals to dust events (Volpe et al., 2009). This indicates that the use of metabolic rates instead of stocks seems more appropriate in tackling the response of biota to atmospheric deposition. Unfortunately, only a couple of studies have so far considered together the changes in metabolic rates induced by atmospheric deposition for both autotrophic and heterotrophic communities: for example, Herut et al. (2005) show that 25 both communities were highly stimulated after fresh dust addition to surface seawater in microcosm experiments with bacterial production (BP) stimulated twice as much as primary production (PP) in the ultra-oligotrophic eastern Mediterranean Sea (increase Introduction of 770 % and 450 %, respectively). Marañon et al. (2010) conducted eight bioassay experiments, over the basin-wide geographical range in the tropical Atlantic and have shown that the effect of Saharan dust deposition on the biological activity of the surface ocean may vary with the degree of oligotrophy of the tested waters. For four out of the eight addition experiments, the relative change in response to dust exceeded 5 a value of 200 % for BP and community respiration (CR), whereas PP never increased by more than 100 % (Marañon et al., 2010). In the western Mediterranean Sea, Pulido-Villena et al. (2008) also show a strong stimulation of bacterial respiration (BR) after dust deposition either in microcosms (125 % average increase in BR) or in situ (100 % increase), the stimulation being proportional to the intensity of the dust flux. Unfortu-10 nately no autotrophic metabolic rate was measured during that study. Depending on the balance between the heterotrophic vs. autotrophic community, the stimulation by atmospheric deposition may thus either result in a CO 2 input or a CO 2 output in oligotrophic environments. Such opposite effects in terms of carbon (C) fate deserve to be explored if we want to understand the role of atmospheric deposition in marine 15 ecosystem functioning, C cycling and feedbacks on climate. Finally, it has been shown recently that aggregation and sorption processes between atmospheric lithogenic particles and in situ (dissolved and particulate) organic matter and subsequent ballasting can induce a strong and rapid particulate organic carbon (POC) export independently of a fertilization effect (Ternon et al., 2010;Bressac and 20 Guieu, 2013;Bressac et al., 2014;Desboeufs et al., 2014). So far, the balance between the different main processes involved in the C cycle after a dust deposition has never been explored. As shown in Guieu et al. (2014b) Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | resentative length of time (7 days after the deposition) is an important improvement compared to the microcosm strategy.
Here we first explore how the balance between bacterial respiration and net primary production evolve after the dust deposition. We then attempt to use the numbers obtained (stocks and fluxes), along with estimates, to quantify how the carbon budget 5 modified by the introduction of dust simulating wet and dry deposition events can be balanced. We report the results for the 3 distinct artificial seeding experiments that were conducted in the frame of the DUNE project in the oligotrophic waters of the Mediterranean Sea in summer 2008 and 2010.

Three seeding experiments during oligotrophic period in the oligotrophic waters of the Mediterranean Sea
The methodology concerning the "large clean mesocosms" (diameter: 2.3 m, surface area: 4.15 m 2 , and volume: 52 m 3 ) setup is fully described in Guieu et al. (2010) and the 2 field campaigns main features are described in a companion paper (Guieu et al.,15 2014b). In this paper we will report on primary production (PP), bacterial respiration (BR) and particulate organic carbon export (POC export ) data acquired during the campaigns DUNE-1 in June 2008 and DUNE-2 in June-July 2010. DUNE-1 consisted in two distinct 8 day experiments: a first simulation of a Saharan wet deposition event (hereafter named "DUNE-P") and a second simulation of a Saharan dry deposition 20 event (hereafter named "DUNE-Q"). DUNE-2 consisted of a single 16 day experiment (hereafter named "DUNE-R") with 2 successive dust wet deposition simulations with 7 days between each seeding (respectively named "R1" and "R2"). Mineral dust deposition was mimicked using the finest fraction (< 20 µm) of alluvial soils collected in a desert soil source area in south Tunisia known to export Aeolian dust to the west-25 ern Mediterranean (Guieu et al., 2010). Dry deposition was mimicked using the raw Introduction soil dispersed in surface seawater whereas wet deposition was mimicked using the same sieved soils after a chemical treatment reproducing cloud water processing by condensation-evaporation. This "evapo-condensed" dust is named hereafter "EC-dust". The methodology to produce those proxies of wet and dry dust deposition is fully described in Guieu et al. (2010). Both dust types had similar chemical composition for 5 nutrients i.e. P, Fe and C, but not for N because of the addition of HNO 3 in the simulated cloud water used to process dust for wet deposition simulation (Guieu et al., 2010). Their respective detailed composition is reported in Desboeufs et al. (2014). For each experiment, 3 mesocosms "CONTROL" without dust addition and 3 mesocosms "DUST" seeded each with a dust flux of 10 g m −2 were deployed. In addition 10 for DUNE-R, a fourth mesocosm was seeded with EC-dust and was devoted to optical measurements (see details in Bressac et al., 2012). The ways the dust was seeded and the mesocosms sampling strategy are fully described in Guieu et al. (2010. Sampling is summarized hereafter. 15 primary productivity (NPP) and particulate organic carbon flux (POC export )

Methodology for sampling and measuring bacteria respiration (BR), net
The methods are described in companion papers of this special issue (see for bacteria respiration: Pulido-Villena et al., 2014, net primary productivity: Ridame et al., 2014and POC export : Bressac et al., 2014, and Desboeufs et al., 2014. The main points are summarized here. The sampling was done every day at the same hours.

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BR -Briefly, BR was determined on 5 m depth samples from each mesocosm gently filtered through a 0.8 µm polycarbonate filter under low vacuum pressure and siphoned into a set of six BOD (biological oxygen demand) bottles. Three BOD bottles from each mesocosm were immediately fixed with Winkler reagents. The other 3 were incubated during 24 h in the dark in a tap water bath to minimize temperature variations and were 25 then fixed with Winkler reagents. Oxygen concentration was measured through a spectrophotometric approach based on measuring the absorbance at 466 nm of the colored I 2 and I − 3 (Labasque et al., 2004, Reinthaler et al., 2006 that the used technique, involving pre-filtration and long incubation time, can lead to an overestimation of the BR rates (see for example, Aranguren-Gassis et al., 2012). This potential overestimation should not be a big problem as far as results are compared with literature data (mostly obtained with the same method) and/or relative changes after dust addition are concerned. We are aware, however, that absolute values of BR 5 or net CO 2 fluxes must be taken with caution. NPP -One sample per depth was collected at both 0.1 and 5 m depth during DUNE-P and -Q and at 5 m depth only during DUNE R, for the determination of NPP using the 13 C uptake determination method. Immediately after sampling, 13 C tracer was added to obtain a final enrichment of about 9 atom% excess. Then, the 13 C-amended bottles 10 were incubated under in situ conditions on a mooring line for 24 h at the corresponding sampling depths. Incubations were ended by filtration onto pre-combusted 25 mm glass fiber filters. Sample filters were stored at −20 • C and dried at 40 • C for 48 h before analysis. Concentration of carbon in particulate matter and 13 C-enrichment were quantified using a ThermoFisher Scientific TM (Bremen, Germany) isotope ratio mass spectrome-15 ter (IRMS) Delta plus, coupled with a C/N analyzer Flash EA via a type III-interface. This net primary productivity corresponds to the net particulate primary productivity. POC export -Samples from sediment traps were preserved using a 5 % buffered solution of formaldehyde. Swimmers were carefully handpicked and the remaining sample was desalted using ultrapure water and freeze-dried. Mass flux was deter-20 mined by weighing the entire freeze-dried sample. The total concentration of carbon (TC) was measured using a CHN analyzer (Perkin Elmer 2400). HNO 3 /HF aciddigestion was performed in 7-mL Teflon flasks at 150 is present both as calcium carbonate and calcium sulfate (Guieu et al., 2010;Desboeufs et al., 2014). The part of Ca associated with sulfate (%Ca CaSO 4 ) was estimated from the particulate S concentration. Therefore, the particulate Ca concentration as carbonate (%Ca CaCO 3 ) corresponded to the difference between total Ca (%Ca) BGD 11,2014 Dust deposition in an oligotrophic marine environment C. Guieu et al. not use the results as unexpected high concentrations and/or variability (either among the 3 depths in a same mesocosm or at the same depth in the triplicate mesocosm were found for many samples, randomly. Unfortunately, the same was observed for filtered samples either transferred in combusted glass ampoules (P and Q experiments) or in acid-washed HDPE bottles (R experiments). This kind of issue with DOC measure-10 ments from samples collected in mesocosms has already been reported and suspected to be attributed to contamination of samples during sample collection and transport as well as during instrument deployment inside the mesocosms (Engel et al., 2013). Only the measurements for in situ ("OUT") measured during DUNE-P will be considered here as a background reference for DOC concentration at the mesocosm site. The seagrasses also characterize our study site in Corsica.

Data integration
The whole data set is presented in Table S1. NPP and BR fluxes were integrated over the mesocosm depth (12.5 m) assuming that the measurement at 5 m is representative of the flux over the entire mesocosm based on the significant similarity (p > 0.05) of the Chl-a concentrations measured at 0.1, 5 and 10 m depth for NPP and on the homogeneity of bacteria abundance for BR (Pulido-Villena, 2013).
BGD 11,2014 Dust deposition in an oligotrophic marine environment C. Guieu et al.

Statistical tests
The mean relative changes (RC, in %) after the P, Q, R1 and R2 seedings were calculated as (X Dust −X Control )×100/(X Dust ) for each parameter (X = NPP, BR and POC export ), where X Control is the mean for the 3 mesocosms CONTROL. In order to compare the results between experiments), means were compared using a one-way ANOVA and a Fisher Least Significant Difference (LSD) means comparison test (α = 0.05). When assumptions for ANOVA were not respected, means were compared using a Kruskal-Wallis test and a post hoc Dunn's test. The statistical tests and the box plots were performed using the Addinsoft TM XLstat software (http://www.xlstat.com).

Orders of magnitude of the background fluxes
The orders of magnitude of all the POC export, NPP and BR fluxes measured in the CONTROL-mescosms and the DUST-mesocosms before seeding for the three experiments are reported in Table 1. Among the 3 parameters, NPP was the more stable flux in the tested waters over the 3 experiments (variation coefficient of 24 %). POC export 15 and BR were more variable but with variation coefficients below 100 % (82 and 68 % respectively). For POC export , no data are reported at such a depth as sediment traps are usually placed below 100 m depth (see synthesis data for the Mediterranean Sea in Ternon et al., 2010). Comparing POC export in surface waters (15 m in the DUNE experi-  Net primary production measured over 7 yr at the DYFAMED site in the Ligurian Sea in the surface mixed layer after the spring period ranged from 2 to 10 µg C L −1 d −1 (Marty and Chiaverini, 2002), which is in good agreement with the NPP rates measured in the In particular, during DUNE-P, BR was higher than previously reported for the open Mediterranean Sea, with values more similar to coastal data as those reported by Navarro et al. (2004). As pointed out by these authors, the production of dissolved organic car-20 bon (DOC) by seagrass meadows, highly abundant in the DUNE study site, may be responsible for such high BR rates. All those comparisons indicate that our study site is a low productive oceanic system with low POC export.
Order of magnitude of induced changes -In order to evaluate the overall impact 25 of the dust deposition on POC export, NPP and BR, the parameters were integrated (POC) or averaged (NPP, BR) over the same duration (7 days) after the artificial seedings. The relative changes (RC in %) compared to CONTROL-Meso are represented in Fig. 1. Average increases relative to CONTROL-meso over 7 days higher than 500 % were observed only for POC export . NPP did not increase after seeding during the Q experiment (dry deposition) and for the other experiments it increased in the range 75-144 %. BR increased in the range 137-156 % in P, Q and R2; a factor 2 higher increase of BR was observed after the R1 seeding (291 %).

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Heterotrophic character of the tested waters -In the literature, the NPP/BR ratio is commonly used to quantify the metabolic status of aquatic systems (see for ex. Del Giorgio et al., 1997;Duarte and Augusti, 1998). If NPP > BR, then the considered system would be a CO 2 sink and on the contrary, if NPP < BR, the ecosystem would be a CO2 source. During DUNE, the average NPP/BR ratio was always ≤ 0.5 in the CONTROL-meso (Fig. 2) indicating that the total carbon processed by bacteria exceeds by far the carbon fixed by phytoplankton: all the tested waters during DUNE before dust addition would thus be a source of dissolved CO 2 . This is in good agreement with recent findings that unproductive aquatic systems are net heterotrophic (del Giorgio et al., 1997). This is reported for the whole Mediterranean sea (Duarte et al.,

A strong heterotrophic character remains after seeding
Although there are some variations in between the triplicate mesocosms as one may expect in natural systems responding to external forcing such as atmospheric deposition, the DUNE data show an increase in both BR and NPP fluxes after seeding (except 5 for NPP for DUNE-Q). The first important point is to note that NPP/BR are below 1 with the average value for the whole data set being 0.32±0.25. As seen on Fig. 3, NPP/BR tends to be significantly lower in DUST-Meso compared to CONTROL-Meso for P and Q indicating that the dust deposition induced an increase in the net heterotrophic character of the tested waters in these experiments.
Indeed, when normalizing to CONTROL values (Fig. 4), there is evidence that the net heterotrophic character of the waters is increasing after seeding for P and Q experiments. Normalized ratio are also on average below 1 after the first seeding of R, but after the second seeding, the ratio is found slightly above 1, signifying that the evolution of the NPP/BR ratio was roughly comparable in both seeded and non-seeded 15 mesocosms. So except for the second seeding of DUNE-R experiment, the dust input did increase the net heterotrophic character of the tested waters.
DUNE setup offers the opportunity to quantify how the carbon budget is perturbed after a simulation by an intense dust deposition onto an oligotrophic system. As discussed in the introduction, until recently most of the studies on impact of dust deposi-20 tion on the surface ocean have focused on the stimulation of phytoplankton. But a first comprehensive study has been published recently indicating how the bacteria community can be stimulated preferentially to autotroph community in the oligotrophic ocean, this stimulation being likely a function of the degree of oligotrophy status where and when the dust deposition does occur (Maranon et al., 2010). Our data are in very good 25 agreement with this previous study as we show that both bacteria activity and primary production have been stimulated by the wet dust input, the stimulation of the bacterial respiration being either higher than that of net CO 2 fixation after the dust event (R1 ex-1719 Introduction

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Interactive Discussion
Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | periment) or equivalent (P and R2). The dust deposition did not thus change drastically the metabolic balance as the tested waters remained net heterotroph. In the 2 cases of DUNE-1 (P, Q), the net heterotrophy was even enhanced by the dust deposition. But this simplistic picture is of course incomplete if other processes that alter the carbon balance are not taken into account.

Carbon balance
As argued in Guieu et al. (2014b), our large clean mesocosms are closed systems with no lateral advection, enclosing a body of water large enough to be representative of the water outside and where stocks and fluxes can be measured after a perturbation such as a simulated dust deposition. Here, we will attempt to use the numbers obtained along with estimates of unmeasured parameters in order to quantify how the carbon budget is affected after dust deposition. The mass balance equation for organic carbon from the mesocosm approach can be written as follows: (δC org /δt) = (δPOC/δt) + (δDOC/δt) = NCP − POC export + At input (1) 15 with NCP, the net community production, POC export , the particulate organic carbon exported downward as measured in the sediment traps, and At input , the amount of organic carbon added by the top to the DUST-meso by seeding (0.31 % of the total dust as determined by Desboeufs et al., 2014).

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NCP is the difference between gross community production (GCP) and community respiration (CR): NCP = GCP − CR. The gross community production can also be expressed by the sum of the net primary production (NPP) that corresponds to the particle primary production in stratified conditions, and of the dissolved primary production (DPP): GCP = NPP + DPP. The community respiration comprises the bacterial respi- although a fraction of the BR can be missed as it was measured on the fraction < 0.8 µm that does not take into account the attached bacteria), CR ≈ 2 BR and Eq. (1) can be rewritten as: (δC org /δt) = (δPOC/δt) + (δDOC/δt) = NPP + DPP − 2BR − POC export + At input (2) 5 Photosynthetic production of dissolved organic carbon (DPP) has been found to be a non-negligible fraction of the total primary production (dissolved and particulate) (see for ex. Baines and Pace, 1991). A similar range of its contribution to total primary production has been found in situ for open ocean oligotrophic systems (i.e 22 % Maranon et al., 2005; 14 %±7 for 9 studies in marine systems reported in Baines and Pace, 1991) and, from a mesocosm study in a coastal productive ecosystem, (19 % Lopez-Sandova et al., 2010). According to recent work in the western Mediterranean Sea in summer (Lagaria et al., 2011), DPP was found to represent ∼ 10 % of the total photosynthetic production. The term DPP in Eq. (2) can thus be replaced by DPP = (0.1· NPP). The calculation of (δC org /δt) is given in Table 2a. Results indicate that the organic 15 carbon sinks, especially respiration, strongly dominate the organic carbon sources, demonstrating that the dust deposition induced a loss of organic carbon of the system in all the experiments, particularly marked after the 3 simulated wet deposition events (P, R). In an attempt to quantify dissolved and particulate partitioning of this organic carbon sink, we have first calculated the changes induced by the seeding inside the meso-20 cosm in the particulate pool (δPOC/δt). (δPOC/δt) is the sum of (1) the changes in the organic carbon pools of phytoplankton, micro-and meso-zooplankton, and bacteria plus (2) a fraction of POC associated to added lithogenic material (POC dust ) (Bressac et al., 2012(Bressac et al., , 2013 and still in suspension inside the mesocosm at the end of the experiment (Desboeufs et al., 2014). Details on the quantification of (δPOC/δt) are given in 25   Tables 2b and S2. Results show that changes in biomass are mostly due to an increase in phytoplankton biomass but that POC dust dominates the particulate organic pool inside the DUST-meso at the end of the experiments (∼ 55-65 % for the 3 wet deposition simulations P and R, and 100 % for Q following the non-stimulation of autotrophs by the dry deposition; Ridame et al., 2014). The remaining term of this carbon budget is the change in the dissolved carbon pool. As emphasized in the methodology section, unfortunately, the non-validated DOC data cannot be used to close this budget. Assuming that the budget is balanced, we 5 deduced the loss in the DOC pool induced by the dust deposition from the available data (Table 2c): we found that the DOC loss was 43 g of C when considering the whole mesocosm, i.e 10 µmol C L −1 d −1 for P experiment, 6 g C mesocosm −1 or On the other hand, as reported by Pulido-Villena et al. (2013) after the 2nd seeding of R experiment, the viral lysis could have fed back in part the pool of DOC. Such mechanism could counterbalance in part the drawdown of DOC necessary to explain the net consumption observed in the absence of allochtonous input of DOC in our setup. Nevertheless, this budget illustrates that in the absence of lateral advection from 5 coastal sources or upwelling from deep waters, such heterotrophic system boosted by atmospheric deposition could decrease significantly the stock of dissolved organic carbon in the surface ocean, a question largely debated at the moment (see for ex. Duarte et al., 2013;Ducklow and Doney, 2013).
Although the net heterotrophy character of the tested waters remained (or even was increased) after the dust addition, POC export was still observed: so the DUNE data also illustrate that a system can be dominated by heterotrophy and still maintain an export of particulate organic carbon even in the absence of allochthonous carbon inputs on the short time scale concerned by the dust deposition event. This is different for open systems that can be heterotrophic while exporting organic carbon vertically only 15 if allochthonous carbon inputs support both fluxes (Cole et al., 2007).

Conclusions
DUNE project allows us the first attempt to evaluate the changes induced in the carbon budget of an oligotrophic system after a simulated Saharan dust deposition above a large body of water during a 7 days period and considering the vertical dimension. 20 We show (i) that the organic carbon sinks dominate the organic carbon sources demonstrating that the dust deposition induced a loss of organic carbon of the system in all the experiments, and (ii) that this loss is due to dissolved organic carbon consumption. Indeed, an overall negative DOC production for the 3 experiments was found. Negative net DOC production indicating net consumption, each of the seeding resulted in a sig- DUNE is an important step forward in the way we understand dust deposition to the ocean in particular because the term "fertilization" is often associated with the a priori belief that dust deposition should increases chlorophyll biomass (despite some evi-5 dences of the contrary from satellite data; Dulac et al., 2004;Volpe et al., 2009) and carbon fixation (and thus increases atmospheric CO 2 drawdown). Instead, DUNE results have shown that by fertilizing predominantly heterotrophic bacteria, dust deposition induces the remineralization of DOC, thereby reducing atmospheric CO 2 drawdown (Rivkin and Anderson, 1997). As emphasized by Pulido-Villena et al., 2008 and this issue, this may reduce substantially the fraction of the dissolved organic carbon stock in the surface ocean that can be exported to deep waters at the time of the winter mixing. Nevertheless, according to the "microbial carbon pump hypothesis" (Jiao et al., 2010), information on the lability of the remaining DOC pool after the enhanced bacterial respiration would be needed to better assess the net effect of dust deposition 15 on DOC export. Other interesting development of this study would be to predict possible scenario for the future using global biogeochemical models where dust pulses such as the ones simulated during DUNE are represented (Guieu et al., 2014a). Indeed on going environmental changes (i.e. increasing temperature and stratification, expansion of the oligotrophic ocean, changes in atmospheric deposition) could modify the role of at-20 mospheric deposition on biogeochemistry of Low-Nutrient-Low-Chlorophyll ocean such as the Mediterranean Sea and an adequate representation of the input of atmospheric new nutrients and particles in those models is timely (Guieu et al., 2014a). In particular, biogeochemical models should consider the important processes highlighted by the DUNE results such as bacteria respiration and (organic matter-Dust) aggregation, Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ran dust: on-board microcosm experiment and field observations, Deep-Sea Res. Pt. II, 52, 3024-3040, 2005. Jiao, N., Herndl, G. J., Hansell, D. A., Benner, R., Kattner, G., Wilhelm, S. W., Kirchman, D. L., Weinbauer, M. G., Luo, T., Chen, F., and Azam, F.: Microbial production of recalcitrant dissolved organic matter: long-term carbon storage in the global ocean, Nat. Rev. Microbiol., 8, Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Navarro, N., Agustí S., and Duarte, C. M.: Plankton metabolism and dissolved organic carbon use in the Bay of Palma, NW Mediterranean Sea, Aquat. Microbial Ecol., 37, 47-54, 2004. Obernosterer, I., Lami, R., Larcher, M., Batailler, N., Catala, P., and Lebaron, P.: Linkage between bacterial carbon processing and the structure of the active bacterial community at a coastal site in the NW Mediterranean Sea, Microb. Ecol., 59, 428-435, 2010. BGD 11,2014 Dust deposition in an oligotrophic marine environment C. Guieu et al.    BGD 11,2014 Dust deposition in an oligotrophic marine environment C. Guieu et al.  BGD 11,2014 Dust deposition in an oligotrophic marine environment C. Guieu et al.