Description: Anthropogenic emissions of carbon dioxide (CO2) and the ongoing accumulation in the surface ocean together with concomitantly decreasing pH and calcium carbonate saturation states have the potential to impact phytoplankton community composition and therefore biogeochemical element cycling on a global scale. Here we report on a recent mesocosm CO2 perturbation study (Raunefjorden, Norway), with a focus on organic matter and phytoplankton dynamics. Cell numbers of three phytoplankton groups were particularly affected by increasing levels of seawater CO2 throughout the entire experiment, with the cyanobacterium Synechococcus and picoeukaryotes (prasinophytes) profiting, and the coccolithophore Emiliania huxleyi (prymnesiophyte) being negatively impacted. Combining these results with other phytoplankton community CO2 experiments into a data-set of global coverage suggests that, whenever CO2 effects are found, prymnesiophyte (especially coccolithophore) abundances are negatively affected, while the opposite holds true for small picoeukaryotes belonging to the class of prasinophytes, or the division of chlorophytes in general. Future reductions in calcium carbonate-producing coccolithophores, providing ballast which accelerates the sinking of particulate organic matter, together with increases in picoeukaryotes, an important component of the microbial loop in the euphotic zone, have the potential to impact marine export production, with feedbacks to Earth's climate system.

Description: The sea surface microlayer (SML) is the thin boundary layer between the ocean and the atmosphere, making it important for air-sea exchange processes. However, little is known about what controls organic matter composition in the SML. In particular, there are only few studies available on the differences of the SML of various oceanic systems. Here, we compared the organic matter and neuston species composition in the SML and the underlying water (ULW) at 11 stations with varying distance from the coast in the Peruvian upwelling regime, a system with high emissions of climate relevant trace gases, such as N2O and CO2. In the open ocean, organic carbon, and amino acids were highly enriched in the SML compared to the ULW. The enrichment decreased at the coastal stations and vanished in the upwelling regime. At the same time, the degradation of organic matter increased from the open ocean to the upwelling stations. This suggests that in the open ocean, upward transport processes or new production of organic matter within the SML are faster than degradation processes. Phytoplankton was generally not enriched in the SML, one group though, the Trichodesmium-like TrL (possibly containing Trichodesmium), were enriched in the open ocean but not in the upwelling region indicating that they find a favorable habitat in the open ocean SML. Our data show that the SML is a distinct habitat; its composition is more similar among different systems than between SML and ULW of a single station. Generally the enrichment of organic matter is assumed to be reduced when encountering low primary production and high wind speeds. However, our study shows the highest enrichments of organic matter in the open ocean which had the lowest primary production and the highest wind speeds.

Description: Climate models project that the Arctic Ocean may experience ice-free summers by the second half of this century. This may have severe repercussions on phytoplankton bloom dynamics and the associated cycling of carbon in surface waters. We currently lack baseline knowledge of the seasonal dynamics of Arctic microbial communities, which is needed in order to better estimate the effects of such changes on ecosystem functioning. Here we present a comparative study of polar summer microbial communities in the ice-free (eastern) and ice-covered (western) hydrographic regimes at the LTER HAUSGARTEN in Fram Strait, the main gateway between the Arctic and North Atlantic Oceans. Based on measured and modeled biogeochemical parameters, we tentatively identified two different ecosystem states (i.e., different phytoplankton bloom stages) in the distinct regions. Using Illumina tag-sequencing, we determined the community composition of both free-living and particle-associated bacteria as well as microbial eukaryotes in the photic layer. Despite substantial horizontal mixing by eddies in Fram Strait, pelagic microbial communities showed distinct differences between the two regimes, with a proposed early spring (pre-bloom) community in the ice-covered western regime (with higher representation of SAR11, SAR202, SAR406 and eukaryotic MALVs) and a community indicative of late summer conditions (post-bloom) in the ice-free eastern regime (with higher representation of Flavobacteria, Gammaproteobacteria and eukaryotic heterotrophs). Co-occurrence networks revealed specific taxon-taxon associations between bacterial and eukaryotic taxa in the two regions. Our results suggest that the predicted changes in sea ice cover and phytoplankton bloom dynamics will have a strong impact on bacterial community dynamics and potentially on biogeochemical cycles in this region.

Description: Gel particles—a class of abundant transparent organic particles—have increasingly gathered attention in marine research. Field studies on the bacterial colonization of marine gels however are still scarce. So far, most studies on respective particles have focused on the upper ocean, while little is known on their occurrence in the deep sea. Here, we report on the vertical distribution of the two most common gel particle types, which are polysaccharide-containing transparent exopolymer particles (TEP) and proteinaceous Coomassie stainable particles (CSP), as well as numbers of bacteria attached to gel particles throughout the water column, from the surface ocean down to the bathypelagial (〈 3,000 m). Our study was conducted in the Arctic Fram Strait during northern hemispheres' summer in 2015. Besides data on the bacterial colonization of the two gel particle types (TEP and CSP), we present bacterial densities on different gel particle size classes according to 12 different sampling depths at four sampling locations. Gel particles were frequently abundant at all sampled depths, and their concentrations decreased from the euphotic zone to the dark ocean. They were colonized by bacteria at all sampled water depths with risen importance at the deepest water layers, where fractions of bacteria attached to gel particles (%) increased within the total bacterial community. Due to the omnipresent bacterial colonization of gel particles at all sampled depths in our study, we presume that euphotic production of this type of organic matter may affect microbial species distribution within the whole water column in the Fram Strait, down to the deep sea. Our results raise the question if changes in the bacterial community composition and functioning on gel particles occur over depth, which may affect microbial respiration and remineralization rates of respective particles in different water layers.

Description: The surface microlayer (SML) is the uppermost thin layer of the ocean and influencing interactions between the air and sea, such as gas exchange, atmospheric deposition and aerosol emission. Organic matter (OM) plays a key role in air-sea exchange processes, but studying how the accumulation of organic compounds in the SML relates to biological processes is impeded in the field by a changing physical environment, in particular wind speed and wave breaking. Here, we studied OM dynamics in the SML under controlled physical conditions in a large annular wind wave channel, filled with natural seawater, over a period of 26 days. Biology in both SML and bulk water was dominated by bacterioneuston and -plankton, respectively, while autotrophic biomass in the two compartments was very low. In general, SML thickness was related to the concentration of dissolved organic carbon (DOC) but not to enrichment of DOC or of specific OM components in the SML. Pronounced changes in OM enrichment and molecular composition were observed in the course of the study and correlated significantly to bacterial abundance. Thereby, hydrolysable amino acids, in particular arginine, were more enriched in the SML than combined carbohydrates. Amino acid composition indicated that less degraded OM accumulated preferentially in the SML. A strong correlation was established between the amount of surfactants coverage and γ-aminobutric acid, suggesting that microbial cycling of amino acids can control physiochemical traits of the SML. Our study shows that accumulation and cycling of OM in the SML can occur independently of recent autotrophic production, indicating a widespread biogenic control of process across the air-sea exchange.

Description: Gel particles, such as transparent exopolymer particles (TEP) and Coomassie stainable particles (CSP), are important organic components in the sea surface microlayer (SML). Here, we present results on the effect of different wind speeds on the accumulation and size distribution of TEP and CSP during a wind wave channel experiment in the Aeolotron. Total areas of TEP (TEPSML) and CSP (CSPSML) in the surface microlayer were exponentially related to wind speed. At wind speeds  〈  6 m s−1, accumulation of TEPSML and CSPSML occurred, decreasing at wind speeds of  〉  8 m s−1. Wind speeds  〉  8 m s−1 also significantly altered the size distribution of TEPSML in the 2–16 µm size range towards smaller sizes. The response of the CSPSML size distribution to wind speed varied through time depending on the biogenic source of gels. Wind speeds  〉  8 m s−1 decreased the slope of CSPSML size distribution significantly in the absence of autotrophic growth. For the slopes of TEP and CSP size distribution in the bulk water, no significant difference was observed between high and low wind speeds. Changes in spectral slopes between high and low wind speed were higher for TEPSML than for CSPSML, indicating that the impact of wind speed on size distribution of gel particles in the SML may be more pronounced for TEP than for CSP, and that CSPSML are less prone to aggregation during the low wind speeds. Addition of an E. huxleyi culture resulted in a higher contribution of submicron gels (0.4–1 µm) in the SML at higher wind speed ( 〉  6 m s−1), indicating that phytoplankton growth may potentially support the emission of submicron gels with sea spray aerosol.

Description: The sea surface microlayer (SML) is located at the air-sea interface, with microorganisms and organic matter in the SML influencing air-sea exchange processes. Yet understanding of the SML bacterial (bacterioneuston) community composition and assembly remains limited. Availability of organic matter, UV radiation and wind speed have previously been suggested to influence the community composition of bacterioneuston. Another mechanism potentially controlling bacterioneuston dynamics is bacterioplankton attached to gel-like particles that ascend through the water column into the SML. We analyzed the bacterial community composition, Transparent Exopolymer Particles (TEP) abundance and nutrient concentrations in the surface waters of the Peruvian upwelling region. The bacterioneuston and bacterioplankton communities were similar, suggesting a close spatial coupling. Four Bacteroidetes families were significantly enriched in the SML, two of them, the Flavobacteriaceae and Cryomorphaceae, were found to comprise the majority of SML-enriched operational taxonomic units (OTUs). The enrichment of these families was controlled by a variety of environmental factors. The SML-enriched bacterial families were negatively correlated with water temperature and wind speed in the SML and positively correlated with nutrient concentrations, salinity and TEP in the underlying water (ULW). The correlations with nutrient concentrations and salinity suggest that the enriched bacterial families were more abundant at the upwelling stations.

Description: The coastal upwelling system off the coast of Peru is characterized by high biological activity and a pronounced subsurface oxygen minimum zone, as well as associated emissions of atmospheric trace gases such as N2O, CH4 and CO2. From 3 to 23 December 2012, R/V Meteor (M91) cruise took place in the Peruvian upwelling system between 4.59 and 15.4° S, and 82.0 to 77.5° W. During M91 we investigated the composition of the sea-surface microlayer (SML), the oceanic uppermost boundary directly subject to high solar radiation, often enriched in specific organic compounds of biological origin like chromophoric dissolved organic matter (CDOM) and marine gels. In the SML, the continuous photochemical and microbial recycling of organic matter may strongly influence gas exchange between marine systems and the atmosphere. We analyzed SML and underlying water (ULW) samples at 38 stations focusing on CDOM spectral characteristics as indicator of photochemical and microbial alteration processes. CDOM composition was characterized by spectral slope (S) values and excitation–emission matrix fluorescence (EEMs), which allow us to track changes in molecular weight (MW) of DOM, and to determine potential DOM sources and sinks. Spectral slope S varied between 0.012 to 0.043 nm−1 and was quite similar between SML and ULW, with no significant differences between the two compartments. Higher S values were observed in the ULW of the southern stations below 15° S. By EEMs, we identified five fluorescent components (F1–5) of the CDOM pool, of which two had excitation/emission characteristics of amino-acid-like fluorophores (F1, F4) and were highly enriched in the SML, with a median ratio SML : ULW of 1.5 for both fluorophores. In the study region, values for CDOM absorption ranged from 0.07 to 1.47 m−1. CDOM was generally highly concentrated in the SML, with a median enrichment with respect to the ULW of 1.2. CDOM composition and changes in spectral slope properties suggested a local microbial release of DOM directly in the SML as a response to light exposure in this extreme environment. In a conceptual model of the sources and modifications of optically active DOM in the SML and underlying seawater (ULW), we describe processes we think may take place (Fig. 1); the production of CDOM of higher MW by microbial release through growth, exudation and lysis in the euphotic zone, includes the identified fluorophores (F1, F2, F3, F4, F5). Specific amino-acid-like fluorophores (F1, F4) accumulate in the SML with respect to the ULW, as photochemistry may enhance microbial CDOM release by (a) photoprotection mechanisms and (b) cell-lysis processes. Microbial and photochemical degradation are potential sinks of the amino-acid-like fluorophores (F1, F4), and potential sources of reworked and more refractory humic-like components (F2, F3, F5). In the highly productive upwelling region along the Peruvian coast, the interplay of microbial and photochemical processes controls the enrichment of amino-acid-like CDOM in the SML. We discuss potential implications for air–sea gas exchange in this area.

Description: Halocarbons are produced naturally in the oceans by biological and chemical processes. They are emitted from surface seawater into the atmosphere, where they take part in numerous chemical processes such as ozone destruction and the oxidation of mercury and dimethyl sulfide. Here we present oceanic and atmospheric halocarbon data for the Peruvian upwelling zone obtained during the M91 cruise onboard the research vessel METEOR in December 2012. Surface waters during the cruise were characterized by moderate concentrations of bromoform (CHBr3) and dibromomethane (CH2Br2) correlating with diatom biomass derived from marker pigment concentrations, which suggests this phytoplankton group is a likely source. Concentrations measured for the iodinated compounds methyl iodide (CH3I) of up to 35.4 pmol L−1, chloroiodomethane (CH2ClI) of up to 58.1 pmol L−1 and diiodomethane (CH2I2) of up to 32.4 pmol L−1 in water samples were much higher than previously reported for the tropical Atlantic upwelling systems. Iodocarbons also correlated with the diatom biomass and even more significantly with dissolved organic matter (DOM) components measured in the surface water. Our results suggest a biological source of these compounds as a significant driving factor for the observed large iodocarbon concentrations. Elevated atmospheric mixing ratios of CH3I (up to 3.2 ppt), CH2ClI (up to 2.5 ppt) and CH2I2 (3.3 ppt) above the upwelling were correlated with seawater concentrations and high sea-to-air fluxes. During the first part of the cruise, the enhanced iodocarbon production in the Peruvian upwelling contributed significantly to tropospheric iodine levels, while this contribution was considerably smaller during the second part.

Description: Responses to ocean acidification in plankton communities were studied during a CO2-enrichment experiment in the Arctic Ocean, accomplished from June to July 2010 in Kongsfjorden, Svalbard (78°56′ 2′′ N, 11°53′ 6′′ E). Enclosed in 9 mesocosms (volume: 43.9–47.6 m3), plankton was exposed to CO2 concentrations, ranging from glacial to projected mid-next-century levels. Fertilization with inorganic nutrients at day 13 of the experiment supported the accumulation of phytoplankton biomass, as indicated by two periods of high chl a concentration. This study tested for CO2 sensitivities in primary production (PP) of particulate organic carbon (PPPOC) and of dissolved organic carbon (PPDOC). Therefore, 14C-bottle incubations (24 h) of mesocosm samples were performed at 1 m depth receiving about 60% of incoming radiation. PP for all mesocosms averaged 8.06 ± 3.64 μmol C L−1 d−1 and was slightly higher than in the outside fjord system. Comparison between mesocosms revealed significantly higher PPPOC at elevated compared to low pCO2 after nutrient addition. PPDOC was significantly higher in CO2-enriched mesocosms before as well as after nutrient addition, suggesting that CO2 had a direct influence on DOC production. DOC concentrations inside the mesocosms increased before nutrient addition and more in high CO2 mesocosms. After addition of nutrients, however, further DOC accumulation was negligible and not significantly different between treatments, indicating rapid utilization of freshly produced DOC. Bacterial biomass production (BP) was coupled to PP in all treatments, indicating that 3.5 ± 1.9% of PP or 21.6 ± 12.5% of PPDOC provided on average sufficient carbon for synthesis of bacterial biomass. During the later course of the bloom, the response of 14C-based PP rates to CO2 enrichment differed from net community production (NCP) rates that were also determined during this mesocosm campaign. We conclude that the enhanced release of labile DOC during autotrophic production at high CO2 exceedingly stimulated activities of heterotrophic microorganisms. As a consequence, increased PP induced less NCP, as suggested earlier for carbon-limited microbial systems in the Arctic.