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Abstract The Mediterranean Sea has been sampled irregularly by research vessels in the past, mostly by national expeditions in regional waters. To monitor the hydrographic, biogeochemical and circulation changes in the Mediterranean Sea, a systematic repeat oceanographic survey programme called Med-SHIP was recommended by the Mediterranean Science Commission (CIESM) in 2011, as part of the Global Ocean Ship-based Hydrographic Investigations Program (GO-SHIP). Med-SHIP consists of zonal and meridional surveys with different frequencies, where comprehensive physical and biogeochemical properties are measured with the highest international standards. The first zonal survey was done in 2011 and repeated in 2018. In addition, a network of meridional (and other key) hydrographic sections were designed: the first cycle of these sections was completed in 2016, with three cruises funded by the EU project EUROFLEETS2. This paper presents the physical and chemical data of the meridional and key transects in the Western and Eastern Mediterranean Sea collected during those cruises.

Biological dinitrogen (N2) fixation is performed solely by specialized bacteria and archaea termed diazotrophs, introducing new reactive nitrogen into aquatic environments. Conventionally, phototrophic cyanobacteria are considered the major diazotrophs in aquatic environments. However, accumulating evidence indicates that diverse non-cyanobacterial diazotrophs (NCDs) inhabit a wide range of aquatic ecosystems, including temperate and polar latitudes, coastal environments and the deep ocean. NCDs are thus suspected to impact global nitrogen cycling decisively, yet their ecological and quantitative importance remain unknown. Here we review recent molecular and biogeochemical evidence demonstrating that pelagic NCDs inhabit and thrive especially on aggregates in diverse aquatic ecosystems. Aggregates are characterized by reduced-oxygen microzones, high C:N ratio (above Redfield) and high availability of labile carbon as compared to the ambient water. We argue that planktonic aggregates are important loci for energetically-expensive N2 fixation by NCDs and propose a conceptual framework for aggregate-associated N2 fixation. Future studies on aggregate-associated diazotrophy, using novel methodological approaches, are encouraged to address the ecological relevance of NCDs for nitrogen cycling in aquatic environments.

Recent studies have indicated that heterotrophic diazotrophs are highly diverse and fix N2 in aquatic environments with potentially adverse conditions for diazotrophy, such as oxic and rich in total nitrogen. In this study, we compared the activity and diversity of heterotrophic diazotrophs associated with aggregates (>12 μm) to free-living cells in the eutrophic Qishon River during the winter and summer seasons. Overall, measured heterotrophic N2 fixation rates in the Qishon River ranged between 2.6–3.5 nmol N L–1 d–1. Heterotrophic N2 fixation was mainly associated with aggregates in the summer samples (74 ± 24%), whereas during the winter the bulk diazotrophic activity was mostly ascribed to the free-living fraction (90 ± 6%). In addition, immunolabeled micrographs indicated the presence of aggregate-associated heterotrophic diazotrophs in both seasons, while phototrophic diazotrophs were also captured during the winter. The richness of free-living and aggregate-associated heterotrophic diazotrophs were overall similar, yet the evenness of the later was significantly smaller, suggesting that few of the species gained advantage from particle lifestyle. The differences in the activity, micro-localization and diversity of the diazotrophic community were mostly attributed to spatiotemporal changes in the ambient C:N ratios (total organic carbon, TOC: total nitrogen) and the TOC concentrations. Taken together, our results shed new light on the contribution of heterotrophic diazotroph associated with aggregates to total heterotrophic N2 fixation in oxic, highly eutrophic aquatic environments.

Nutrient pollution may negatively affect the water quality and ecological status of rivers and estuaries worldwide, specifically in stratified and small inland estuaries. We present a long-term, two-decade data set of dissolved inorganic nutrient concentrations, chlorophyll-a (chl-a), dissolved oxygen (DO), and potentially toxic algal cell concentrations at the Kishon River estuary (Israel) as a case study for assessing nutrient ecological thresholds in such type of estuaries, prevalent along the Mediterranean coast of Israel. In-situ measurements and water samples were collected at 3 permanent stations at the lower part of the estuary every March and October/November in 40 campaigns over the years 2002 to 2021. In spite of an improvement in nutrient loads and concentrations as recorded over the last 2 decades, the nutrient and chl-a levels at the Kishon estuary surface water represent mostly a ‘bad’ or ‘moderate’ ecological state, considering the recommended thresholds discussed in this study. It is suggested to develop a combined suite of nutrient and biological variables for assessing Good Environmental Status (GES), considering the relatively high residence time of such small, low-flow estuarine water bodies.

Long-term trends in oxygen, salinity, and nutrients were followed in the Southeastern Mediterranean (SEMS) deep waters from 2002 to 2020. Results show a net decrease in oxygen since 2008 of −0.5 ± 0.1 μmol kg−1 yr−1 in the bathypelagic depths (1,200–2,000 m). Multiannual variability in oxygen levels superimposed this trend, and is likely associated with variations in thermohaline fluxes. The 2020 mean oxygen concentration of 179.5 ± 2.3 μmol kg−1 is comparable to the pre-Eastern Mediterranean Transient (EMT) mean value. The post-EMT signature is clearly demonstrated in both oxygen and salinity over the period of 2002–2013, but since 2014 it diminished, mainly due to mixing of the Aegean deep water (AegDW) mass with the overlying old Adriatic water mass. This trend reflects a switch back to the pre-EMT regime, characterized by thermohaline homogeneity of the deep water column in the SEMS. The long-term decline of deep water oxygen levels is also accompanied by a corresponding increase in dissolved inorganic nutrients, supporting aging of the deep water masses. Our results suggest that ventilation of the SEMS deep water is currently occurring at a lower, pre-EMT rate, probably as a result of moderated deep water formation in recent time.

Submarine groundwater discharge (SGD) has been shown to be an important source of nutrients in coastal environments, especially nitrogen and silica, and thereby relive nutrient limitation to phytoplankton. Here, we followed autotrophic microbial biomass, activity, and community composition at a site strongly influenced by SGD and a nearby nutrients-poor reference site at the oligotrophic Israeli shallow rocky coast [southeastern Mediterranean Sea (SEMS)] between 2011 and 2019. The surface water at the SGD-affected area had significantly higher NO3 + NO2 (∼10-fold) and Si(OH)4 (∼2-fold) levels compared to the reference site, while no significant differences were observed for PO4 or NH4. This resulted in a significant increase in algae biomass (∼3.5-fold), which was attributed to elevated Synechococcus (∼3.5-fold) and picoeukaryotes (∼2-fold) at the SGD-affected site, and in elevated primary production rates (∼2.5-fold). Contrary to most SGD-affected coastal areas, diatoms biomass remained unchanged between sites, despite the elevated N and Si, suggesting the dominance of picophytoplankton over microphytoplankton at the SEMS. DNA sequencing of the 16S and 18S rDNA supported these findings. These results highlight the influence of SGD on shallow-water microbial populations. Our observations are consistent with recent studies showing that phytoplankton along the Israeli coast are likely nitrogen + silica limited, and may have important ecological and regulatory implications for environmental policy and management of coastal aquifers.

Planktonic heterotrophic diazotrophs (N2-fixers) are widely distributed in marine and freshwater systems, yet limited information is available on their activity, especially in environments with adverse conditions for diazotrophy (e.g., N-rich and oxygenated). Here, we followed the localization and activity of heterotrophic diazotrophs in the hyper-eutrophic N-rich Qishon River—an environment previously considered to be unfavorable for diazotrophy. Our results indicate high heterotrophic N2 fixation rates (up to 6.9 nmol N L–1 d–1), which were approximately three fold higher at an upstream location (freshwater) compared to an estuary (brackish) site. Further, active heterotrophic diazotrophs were capture associated with free-floating aggregates by a newly developed immunolocalization approach. These findings provide new insights on the activity of heterotrophic diazotrophs on aggregates in environments previously considered with adverse conditions for diazotrophy. Moreover, these new insights may be applicable to other aquatic regimes worldwide with similar N-rich/oxygenated conditions that should potentially inhibit N2 fixation.

Abstract The southeastern Mediterranean Sea (SEMS) is a warm and sunlit marine environment with low ambient N concentration, thus considered ideal for diazotrophy by autotrophic diazotrophs such as Trichodesmium. Despite the favorable conditions, N2 fixation rates are often low and Trichodesmium has hardly been spotted in the SEMS. This study reports on the occurrence of a Trichodesmium bloom in the SEMS which was ascribed to T. erythraeum according to DNA fingerprinting of the nifH gene. We found that this bloom (1407 ± 983 cells L−1) was triggered by an intense outburst of raw sewage that supplied high concentrations of N, P and dissolved organic carbon (DOC), which resulted in low N:P (~12:1) and exceptionally high C:P (~1340:1) ratios. We surmise that these conditions provided favorable conditions for Trichodesmium bloom to form via mixotrophic metabolism. As a result, a fourfold increase in N2 fixation was recorded, which contributed ~70% to new primary production and spur a sharp increase in phytoplankton activity and biomass. The conclusions of this study point on a new paradigm for bloom-forming T. erythraeum which is tightly linked to anthropogenic sources and prompt microbial productivity in oligotrophic marine environments such as the SEMS.

Phytoplankton and heterotrophic bacteria rely on a suite of inorganic and organic macronutrients to satisfy their cellular needs. Here, we explored the effect of dissolved inorganic phosphate (PO4) and several dissolved organic molecules containing phosphorus [ATP, glucose-6-phosphate, 2-aminoethylphosphonic acid, collectively referred to as dissolved organic phosphorus (DOP)], on the activity and biomass of autotrophic and heterotrophic microbial populations in the coastal water of the southeastern Mediterranean Sea (SEMS) during summertime. To this end, surface waters were supplemented with PO4, one of the different organic molecules, or PO4 + ATP, and measured the PO4 turnover time (Tt), alkaline phosphatase activity (APA), heterotrophic bacterial production (BP), primary production (PP), and the abundance of the different microbial components. Our results show that PO4 alone does not stimulate any significant change in most of the autotrophic or heterotrophic bacterial variables tested. ATP addition (alone or with PO4) triggers the strongest increase in primary and bacterial productivity or biomass. Heterotrophic bacterial abundance and BP respond faster than phytoplankton (24 h post addition) to the various additions of DOP or PO4 + ATP, followed by a recovery of primary productivity (48 h post addition). These observations suggest that both autotrophic and heterotrophic microbial communities compete for labile organic molecules containing P, such as ATP, to satisfy their cellular needs. It also suggests that SEMS coastal water heterotrophic bacteria are likely C and P co-limited.

The marine cyanobacterium Prochlorococcus is a dominant photoautotroph in many oligotrophic Low-Nutrients-Low-Chlorophyll (LNLC) regions. While the chemical impact of aerosols upon interaction with surface seawater was documented in numerous studies, we show that Prochlorococcus cells are affected also by bio-aerosols (potentially biological agents in the dust/aerosols such as membrane-bound extracellular vesicles, small-size bacteria and/or viruses), resulting in lower surface seawater abundances in the oligotrophic Mediterranean Sea. We conducted experimental amendments of ‘live’ aerosol/dust particles and aerosol filtrates (Prochlorococcus cultures (MED4). Results show a significant decline in cell biomass (Prochlorococcus. Accordingly, the dominance of Synechococcus over Prochlorococcus throughout the surface Mediterranean Sea (observed mainly in spring when atmospheric aerosol levels are relatively high) and the lack of spatial westward gradient in Prochlorococcus biomass as typically observed for chlorophyll-a or other cyanobacteria may be attributed, at least to some extent, to the impact of bio-aerosol deposition across the basin. Predictions for enhanced desertification and increased dust emissions may intensify the transport and potential impact of bio-aerosols in LNLC marine systems.

Mesopelagic prokaryotes (archaea and bacteria), which are transported together with nutrient-rich intermediate-water to the surface layer by deep convection in the oceans (e.g., winter mixing, upwelling systems), can interact with surface microbial populations. This interaction can potentially affect production rates and biomass of surface microbial populations, and thus play an important role in the marine carbon cycle and oceanic carbon sequestration. The Eastern Mediterranean Sea (EMS) is one of the most oligotrophic and warm systems in the world's oceans, with usually very shallow winter mixing (

The effects of atmospheric deposition on plankton community structure were examined during a mesocosm experiment using water from the Cretan Sea (Eastern Mediterranean), an area with a high frequency of atmospheric aerosol deposition events. The experiment was carried out under spring-summer conditions (May 2012). The main objective was to study the changes induced from a single deposition event, on the autotrophic and heterotrophic surface microbial populations, from viruses to zooplankton. To this end, the effects of Saharan dust addition were compared to the effects of mixed aerosol deposition on the plankton community over 9 days. The effects of the dust addition seemed to propagate throughout the food-web, with changes observed in nearly all of the measured parameters up to copepods. The dust input stimulated increased productivity, both bacterial and primary. Picoplankton, both autotrophic and heterotrophic capitalized on the changes in nutrient availability and microzooplankton abundance also increased due to increased availability of prey. Five days after the simulated deposition, copepods also responded, with an increase in egg production. The results suggest that nutrients were transported up the food web through autotrophs, which were favored by the Nitrogen supplied through both treatments. Although, the effects of individual events are generally short lived, increased deposition frequency and magnitude of events is expected in the area, due to predicted reduction in rainfall and increase in temperature, which can lead to more persistent changes in plankton community structure. Here we demonstrate how a single dust deposition event leads to enhancement of phytoplankton and microzooplankton and can eventually, through copepods, transport more nutrients up the food web in the Eastern Mediterranean Sea.

Prokaryotic microbes can become aerosolized and deposited into new environments located thousands of kilometers away from their place of origin. The Mediterranean Sea is an oligotrophic to ultra-oligotrophic marginal sea, which neighbors northern Africa (a major source of natural aerosols) and Europe (a source of mostly anthropogenic aerosols). Previous studies demonstrated that airborne bacteria deposited during dust events over the Mediterranean Sea may significantly alter the ecology and function of the surface seawater layer, yet little is known about their abundance and diversity during ‘background’ non-storm conditions. Here, we describe the abundance and genetic diversity of airborne bacteria in 16 air samples collected over an East-West transect of the entire Mediterranean Sea during non-storm conditions in April 2011. The results show that airborne bacteria represent diverse groups with the most abundant bacteria from the Firmicutes (Bacilli and Clostridia) and Proteobacteria (Alphaproteobacteria, Betaproteobacteria, and Gammaproteobacteria) phyla. Most of the bacteria in our samples have previously been observed in the air at other open ocean locations, in the air over the Mediterranean Sea during dust storms, and in the Mediterranean seawater. Airborne bacterial abundance ranged from 0.7 × 104 to 2.5 × 104 cells m−3 air, similar to abundances at other oceanic regimes. Our results demonstrate that airborne bacterial diversity is positively correlated with the mineral dust content in the aerosols and was spatially separated between major basins of the Mediterranean Sea. To our knowledge, this is the first comprehensive biogeographical dataset to assess the diversity and abundance of airborne microbes over the Mediterranean Sea. Our results shed light on the spatiotemporal distribution of airborne microbes and may have implications for dispersal and distribution of microbes (biogeography) in the ocean.

The northern Red Sea (NRS) is a low-nutrient, low-chlorophyll (LNLC) ecosystem with high rates of atmospheric deposition due to its proximity to arid regions. Impacts of atmospheric deposition on LNLC ecosystems have been attributed to the chemical constituents of dust, while overlooking bioaerosols. Understanding how these vast areas of the ocean will respond to future climate and anthropogenic change hinges on the response of microbial communities to these changes. We tested the impacts of bioaerosols on the surface water microbial diversity and the primary and bacterial production rates in the NRS, a system representative of other LNLC oceanic regions, using a mesocosm bioassay experiment. By treating NRS surface seawater with dust, which contained nutrients, metals, and viable organisms, and “UV-treated dust” (which contained only nutrients and metals), we were able to assess the impacts of bioaerosols on local natural microbial populations. Following amendments (20 and 44 h) the incubations treated with “live dust” showed different responses than those with UV-treated dust. After 44 h, primary production was suppressed (as much as 50%), and bacterial production increased (as much as 55%) in the live dust treatments relative to incubations amended with UV-treated dust or the control. The diversity of eukaryotes was lower in treatments with airborne microbes. These results suggest that the airborne microorganisms and viruses alter the surface microbial ecology of the NRS. These results may have implications for the carbon cycle in LNLC ecosystems, which are expanding and are especially important since dust storms are predicted to increase in the future due to desertification and expansion of arid regions.

Airborne prokaryotes are transported along with dust/aerosols, yet very little attention is given to their temporal variability above the oceans and the factors that govern their abundance. We analyzed the abundance of autotrophic (cyanobacteria) and heterotopic airborne microbes in 34 sampling events between 2015−2018 at a coastal site in the SE Mediterranean Sea. We show that airborne autotrophic (0.2−7.6 cells × 103 m−3) and heterotrophic (0.2−30.6 cells × 103 m−3) abundances were affected by the origin and air mass trajectory, and the concentration of dust/aerosols in the air, while seasonality was not coherent. The averaged ratio between heterotrophic and autotrophic prokaryotes in marine-dominated trajectories was ~1.7 ± 0.6, significantly lower than for terrestrial routes (6.8 ± 6.1). Airborne prokaryotic abundances were linearly and positively correlated to the concentrations of total aerosol, while negatively correlated with the aerosol’s anthropogenic fraction (using Pb/Al or Cu/Al ratios as proxies). While aerosols may play a major role in dispersing terrestrial and marine airborne microbes in the SE Mediterranean Sea, the mechanisms involved in the dispersal and diversity of airborne microorganisms remain to be studied and should include standardization in collection and analysis protocols.

Recent estimates of nutrient budgets for the Eastern Mediterranean Sea (EMS) indicate that atmospheric aerosols play a significant role as suppliers of macro- and micro- nutrients to its Low Nutrient Low Chlorophyll water. Here we present the first mesocosm experimental study that examines the overall response of the oligotrophic EMS surface mixed layer (Cretan Sea, May 2012) to two different types of natural aerosol additions, pure Saharan dust (SD, 1.6 mg l-1) and mixed aerosols (A - polluted and desert origin, 1 mg l-1). We describe the rationale, the experimental set-up, the chemical characteristics of the ambient water and aerosols and the relative maximal biological impacts that resulted from the added aerosols. The two treatments, run in triplicates (3 m3 each), were compared to control-unamended runs. Leaching of approximately 2.1-2.8 and 2.2-3.7 nmol PO4 and 20-26 and 53-55 nmol NOx was measured per each milligram of SD and A, respectively, representing an addition of approximately 30% of the ambient phosphate concentrations. The nitrate/phosphate ratios added in the A treatment were twice than those added in the SD treatment. Both types of dry aerosols triggered a positive change (25-600% normalized per 1 mg l-1 addition) in most of the rate and state variables that were measured: bacterial abundance (BA), bacterial production (BP), Synechococcus (Syn) abundance, chlorophyll-a (chl-a), primary production (PP) and dinitrogen fixation (N2-fix), with relative changes among them following the sequence BP>PP≈N2-fix>chl-a≈BA≈Syn. Our results show that the ‘polluted’ aerosols triggered a relatively larger biological change compared to the SD amendments (per a similar amount of mass addition), especially regarding BP and PP. We speculate that despite the co-limitation of P and N in the EMS, the additional N released by the A treatment may have triggered the relatively larger response in most of the rate and state variables as compared to SD. An implication of our study is that a warmer atmosphere in the future may increase dust emissions and influence the intensity and length of the already well stratified water column in the EMS and hence the impact of the aerosols as a significant external source of new nutrients.

The oligotrophic southeastern Mediterranean Sea (SEMS) is frequently exposed to desert-dust deposition which supplies nutrients, trace metals and a wide array of viable airborne microorganisms. In this study, we experimentally examined the impact of aerosol addition, collected during an intense dust storm event in early September 2015, on the biomass and activity of pico-phytoplankton and heterotrophic bacterial populations at the sea-surface micro layer (SML) relative to the sub surface layer (SSL). Aerosol (1.5 mg L-1) was added to SML and SSL water samples in microcosms (4.5 L) and the water was frequently sampled over a period of 48 hours. While the aerosol amendment triggered a moderate 1.5-2 fold increase in primary production in both the SML and the SSL, bacterial production increased by ~3 and ~7 folds in the SSL and SML, respectively. Concurrently, the abundance and flow-cytometric characteristics (green fluorescence and side scatter signals) of high nucleic acid (HNA) and low nucleic acid (LNA) bacterial cells showed a significant increase in the %HNA, in both SML and SSL samples following aerosol amendment. This shift in nucleic acid content took place at a much faster rate in the SML, suggesting a more active heterotrophic community. These changes were likely a result of higher rates of carbon utilizations in the SML following the dust addition, as assessed by a selected hydrocarbons and saccharides analysis. Additionally, a high absorption rate of hydrocarbons by the aerosol particles was measured following the additions, leaving less than 10% of these molecules available for potential heterotrophic microbial utilization. Our results suggest that the heterotrophic microbial community inhabiting the SML is more efficient in utilizing aerosol associated constituents than the community in the SSL.

The atmospheric deposition of gases and particulates from the Sahara Desert and European landmass is an important source of nutrients for the Mediterranean Sea. In this study, we investigated how such atmospheric input might affect bacterial metabolic activities and community dynamics in the ultra-oligotrophic Eastern Mediterranean Sea. Thus a mesocosm simulation experiment was conducted using pure Saharan dust (SD) and mixed aerosols (A, polluted and desert origin). The cell specific bacterial production (BP) was stimulated soon after the addition of SD and A, with a higher degree of stimulation being observed in the activity of Alphaproteobacteria than in Gammaproteobacteria, and this lead to significant increases in community BP. Subsequently, a shift between these two dominating classes was observed (such that the proportion of Gammaproteobacteria increased while that of Alphaproteobacteria decreased), along with significant increases in bacterial abundance and chlorophyll a concentration. After a few days, although the abundance of bacteria was still significantly higher in the SD- or A-treated groups, differences in the active community composition between the treatment and control groups were reduced. The altered activity of the two dominating Proteobacteria classes observed, might reflect their different strategies in responding to external nutrient input: with Alphaproteobacteria being more responsive to the direct dust input, whereas Gammaproteobacteria seemed to benefit more from the increase in phytoplankton biomass. In addition, the input of A had a stronger immediate effect and longer lasting influence on changing the active bacterial community composition than did that of SD. Our findings show that episodic atmospheric deposition events might affect the microbial community with regards to their abundance, activity and composition over a short period of time, and thus regulate the function of the microbial community and carbon cycling in oligotrophic waters.

Large amounts of dust and atmospheric aerosols, originating from surrounding desert areas (e.g., Sahara and Middle East) are deposited annually on the surface of the Eastern Mediterranean Sea. These depositions can provide high amounts of micro (such as Fe, Zn, Co) and macro nutrients (such as P and N) to supplement nutrient-poor surface waters- that typically limit primary productivity and also dinitrogen (N2) fixation in many marine environments. Here, we studied the impact of the atmospheric deposition of dust and aerosols on N2 fixation in the Cretan Sea (Eastern Mediterranean Sea). Mixed polluted aerosols (hereafter A) and Saharan dust (hereafter SD) were added to nine mesocosms (3-m3 each) containing surface mixed layer seawater (~10 m), and N2 fixation was evaluated for 6 days during May 2012 (springtime). The addition of SD triggered a rapid (30 h) and robust (2-4 fold) increase in N2 fixation rates that remained high for 6 days and contributed 3-8% of the primary productivity. The A addition also resulted in higher N2 fixation rates compared to the unamended control mesocosms, although the responses were less profound (1.5-2 fold) and accounted for only 2-4% of the primary productivity. The microbial community responded differently to the two additions. Heterotrophic bacterial N2 fixers dominated the diazotroph community in A and the control mesocosms, while the non-filamentous cyanobacterial group Trichodesmium prevailed in the SD treatment (68% of all the operational taxonomic units, verified by qPCR analyses). Our results indicate that the aerosol source, its route prior to deposition, and its specific chemical composition, can alter the diazotrophic diversity and activity in the Eastern Mediterranean Sea and may thus impact both the N and C dynamics in this impoverished environment.

Acid processes in the atmosphere, particularly those caused by anthropogenic acid gases, increase the amount of bioavailable P in dust and hence are predicted to increase microbial biomass and primary productivity when supplied to oceanic surface waters. This is likely to be particularly important in the Eastern Mediterranean Sea (EMS), which is P limited during the winter bloom and N&P co-limited for phytoplankton in summer. However, it is not clear how the acid processes acting on Saharan dust will affect the microbial biomass and primary productivity in the EMS. Here, we carried out bioassay manipulations on EMS surface water on which Saharan dust was added as dust (Z), acid treated dust (ZA), dust plus excess N (ZN) and acid treated dust with excess N (ZNA) during springtime (May 2012) and measured bacterioplankton biomass, metabolic and other relevant chemical and biological parameters. We show that acid treatment of Saharan dust increased the amount of bioavailable P supplied by a factor of ~40 compared to non-acidified dust (18.4 nmoles P mg-1 dust vs. 0.45 nmoles P mg-1 dust, respectively). The increase in chlorophyll, primary and bacterial productivity for treatments Z and ZA were controlled by the amount of N added with the dust while those for treatments ZN and ZNA (in which excessive N was added) were controlled by the amount of P added. These results confirm that the surface waters were N&P co-limited for phytoplankton during springtime. However, total chlorophyll and primary productivity in the acid treated dust additions (ZA and ZNA) were less than predicted from that calculated from the amount of the potentially limiting nutrient added. This biological inhibition was interpreted as being due to labile trace metals being added with the acidified dust. A probable cause for this biological inhibition was the addition of dissolved Al, which forms potentially toxic Al nanoparticles when added to seawater. Thus, the effect of anthropogenic acid processes in the atmosphere, while increasing the flux of bioavailable P from dust to the surface ocean, may also add toxic trace metals such as Al, which moderate the fertilizing effect of the added nutrients.

This study explores the potential impacts of microbes deposited into the surface seawater of the southeastern Mediterranean Sea (SEMS) along with atmospheric particles on marine autotrophic and heterotrophic production. We compared in situ changes in autotrophic and heterotrophic microbial abundance and production rates before and during an intense dust storm event in early September 2015. Additionally, we measured the activity of microbes associated with atmospheric dry deposition (also referred to as airborne microbes) in sterile SEMS water using the same particles collected during the dust storm. A high diversity of prokaryotes and a low diversity of autotrophic eukaryotic algae were delivered to surface SEMS waters by the storm. Autotrophic airborne microbial abundance and activity were low, contributing ~1% of natural abundance in SEMS water and accounting for 1-4% to primary production. Airborne heterotrophic bacteria comprised 30-50% of the cells and accounted for 13-42% of bacterial production. Our results demonstrate that atmospheric dry deposition may supply not only chemical constitutes but also microbes that can affect ambient microbial populations and their activity in the surface ocean. Airborne microbes may play a greater role in ocean biogeochemistry in the future in light of the expected enhancement of dust storm durations and frequencies due to climate change and desertification processes.

Atmospheric dust/aerosol deposition is an important source of external nutrients for the surface of the ocean. This study shows high-resolution observational data gathered in situ over a period of four years on bacterial and phytoplankton abundance and activity during typical background atmospheric conditions and during intense dust storm events in the low-nutrient, low-chlorophyll (LNLC) coastal waters of the southeastern Mediterranean Sea (SEMS). Chlorophyll a (an estimate for phytoplankton biomass) and bacterial abundance show moderate changes in response to dust deposition/events (−10% and +20%, respectively), while primary production, bacterial production, and N2 fixation rates were all significantly and positively affected by deposition (+25 to +40%; p < 0.05). The rapid changes in bacterial and/or phytoplankton rate parameters suggest that the released micro-/macronutrients from atmospheric deposition are tunneled directly in metabolic processes and, to a lesser extent, for biomass accumulation. The predicted expansion of LNLC areas in oceans in the future, and the projected increase in dust emission due to desertification, may affect the production of marine microbial communities in the surface of the ocean, yet only moderately affect their biomass or standing stock. Such alterations may impact carbon sequestration to the deep ocean.

This study considers variability in phytoplankton and heterotrophic bacterial abundances and production rates, in one of the most oligotrophic marine regions in the world-the Levantine Basin. The temporal dynamics of these planktonic groups were studied in the coastal waters of the southeastern Mediterranean Sea approximately every two weeks for a total of two years. Heterotrophic bacteria were abundant mostly during late summer and midwinter, and were positively correlated with bacterial production and with N2 fixation. Based on size fractionating, picophytoplankton was abundant during the summer, whereas nano-microphytoplankton predominated during the winter and early spring, which were also evident in the size-fractionated primary production rates. Autotrophic abundance and production correlated negatively with temperature, but did not correlate with inorganic nutrients. Furthermore, a comparison of our results with results from the open Levantine Basin demonstrates that autotrophic and heterotrophic production, as well as N2 fixation rates, are considerably higher in the coastal habitat than in the open sea, while nutrient levels or cell abundance are not different. These findings have important ecological implications for food web dynamics and for biological carbon sequestration in this understudied region.