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: Highlights: • Activity of Arctic bacterioplankton in summer is regulated by concentration and composition of organic matter. • Bacterial production in Fram Strait is significantly related to concentrations of total amino acids. • Bacterioplankton in Polar Water show enhanced enzymatic hydrolysis of combined carbohydrates compared to Atlantic Water. Abstract The bacterial turnover of organic matter was investigated in Fram Strait at 79°N. Both Atlantic Water (AW) inflow and exported Polar Water (PW) were sampled along a transect from Spitsbergen to the eastern Greenland shelf during a late successional stage of the main annual phytoplankton bloom in summer. AW showed higher concentrations of amino acids than PW, while organic matter in PW was enriched in combined carbohydrates. Bacterial growth and degradation activity in AW and PW were related to compositional differences of organic matter. Bacterial production and leucine-aminopeptidase along the transect were significantly correlated with concentrations of amino acids. Activity ratios between the extracellular enzymes β-glucosidase and leucine-aminopeptidase indicate the hydrolysis potential for polysaccharides relative to proteins. Along the transect, these ratios showed a higher hydrolysis potential for polysaccharides relative to proteins in PW than in AW, thus reflecting the differences in organic matter composition between the water masses. Q10 values for bacterial production ranged from 2.4 (± 0.8) to 6.0 (± 6.8), while those for extracellular enzymes showed a broader range of 1.5 (± 0.5) to 23.3 (± 11.8). Our results show that in addition to low seawater temperature also organic matter availability contributes to the regulation of bacterial growth and enzymatic activity in the Arctic Ocean.

Description: Two cruises were conducted after the diatom spring bloom in the northern Bay of Biscay (2006, 2007), to assess the contribution of combined carbohydrates to organic carbon partitioning. Partitioning of total organic carbon (TOC) into particulate organic carbon (POC) and dissolved organic carbon (DOC) differed between the two years, particularly for depths above 60 m, and was related to the vernal development of the system: a post spring-bloom system in 2007, and a more stratified summer system with higher coccolithophore abundance in 2006. In general, contribution of POC to TOC ranged between 4% and 28% and decreased with depth. Concentration of high-molecular-weight (HMW;〉1 kDa) dissolved combined carbohydrates (dCCHO) ranged from 0.6 to 1.4 μmol L−1 and contributed between 4% and 11% to DOC. Concentration of particulate combined carbohydrates (pCCHO) varied between 0.03 and 1.3 μmol L−1. A high contribution of pCCHO to POC was observed in 2007, i.e. 22–60% C compared to 3–10% C in 2006, and coincided with a higher abundance of transparent exopolymer particles (TEP). TEP accounted for 0.4–2.0 μmol C L−1 in 2007 and 0.5–1.5 μmol C L−1 in 2006. Above 60 m, differences in contribution of TEP-C to POC were most pronounced yielding 15.4±3.0% in 2007 compared to relatively low 4.8±1.4%, in 2006. TEP-C could explain about 60% in 2007 and about 40% of pCCHO-C in 2006. Hence, TEP were identified as a substantial component of pCCHO and POC, particularly in the wake of the spring bloom. Molecular composition of CCHO, i.e. HMW−dCCHO+pCCHO, revealed little difference between the years but strong variation over depth. Uronic acids (URA) were identified as a major component of CCHO (20–40%). Our study indicates that the distribution and composition of CCHO in surface seawater are determined by biogeochemical processes on a seasonal scale. A better knowledge of CCHO cycling and molecular signature has therefore a high potential for a better tracing of carbon dynamics in shelf sea ecosystems. Highlights: ► Role of combined carbohydrates for DOC–POC partitioning assessed (Bay of Biscay). ► TEP comprised substantial carbon fraction of particulate carbohydrates. ► Molecular composition of polysaccharides revealed organic matter diagenesis. ► Uronic acids comprised 20–40% of total and dissolved combined carbohydrates.

Description: The distribution of transparent exopolymer particles (TEP) was investigated during a coccolithophorid bloom in the northern Bay of Biscay (North Atlantic Ocean) in early June 2006. MODIS chlorophyll-a (Chl-a) and reflectance images before and during the cruise were used to localize areas of important biological activity and high reflectance (HR). TEP profiles along the continental margin, determined using microscopic (TEPmicro) and colorimetric (TEPcolor) methods, showed abundant (6.1×106–4.4×107 L−1) and relatively small (0.5–20 μm) particles, leading to a low total volume fraction (0.05–2.2 ppm) of TEPmicro and similar vertical profiles of TEPcolor. Estimates of carbon content in TEP (TEP-C) derived from the microscopic approach yielded surface concentration of 1.50 μmol C L−1. The contribution of TEP-C to particulate organic carbon (POC) was estimated to be 12% (molar C ratio) during this survey. Our results suggest that TEP formation is a probable first step to rapid and efficient export of C during declining coccolithophorid blooms.

Description: Primary production (PP), calcification (CAL), bacterial production (BP) and dark community respiration (DCR) were measured along with a set of various biogeochemical variables, in early June 2006, at several stations at the shelf break of the northern Bay of Biscay. The cruise was carried out after the main spring diatom bloom that, based on the analysis of a time-series of remotely sensed chlorophyll-a (Chl-a), peaked in mid-April. Remotely sensed sea surface temperature (SST) indicated the occurrence of enhanced vertical mixing (due to internal tides) at the continental slope, while adjacent waters on the continental shelf were stratified, as confirmed by vertical profiles of temperature acquired during the cruise. The surface layer of the stratified water masses (on the continental shelf) was depleted of inorganic nutrients. Dissolved silicate (DSi) levels probably did not allow significant diatom development. We hypothesize that mixing at the continental slope allowed the injection of inorganic nutrients that triggered the blooming of mixed phytoplanktonic communities dominated by coccolithophores (Emiliania huxleyi) that were favoured with regards to diatoms due to the low DSi levels. Based on this conceptual frame, we used an indicator of vertical stratification to classify the different sampled stations, and to reconstruct the possible evolution of the bloom from the onset at the continental slope (triggered by vertical mixing) through its development as the water mass was advected on-shelf and stratified. We also established a carbon mass balance at each station by integrating in the photic layer PP, CAL and DCR. This allowed computation at each station of the contribution of PP, CAL and DCR to CO2 fluxes in the photic layer, and how they changed from one station to another along the sequence of bloom development (as traced by the stratification indicator). This also showed a shift from net autotrophy to net heterotrophy as the water mass aged (stratified), and suggested the importance of extracellular production of carbon to sustain the bacterial demand in the photic and aphotic layers.

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

Description: The Arctic Ocean plays a key role in regulating the global climate, while being highly sensitive to climate change. Temperature in the Arctic increases faster than the global average, causing a loss of multiyear sea-ice and affecting marine ecosystem structure and functioning. As a result, Arctic primary production and biogeochemical cycling are changing. Here, we investigated inter-annual changes in the concentrations of particulate and dissolved organic carbon (POC, DOC) together with biological drivers, such as phyto- and bacterioplankton abundance in the Fram Strait, the Atlantic gateway to the Central Arctic Ocean. Data have been collected in summer at the Long-Term Ecological Research observatory HAUSGARTEN during eight cruises from 2009 to 2017. Our results suggest that the dynamic physical system of the Fram Strait induces strong heterogeneity of the ecosystem that displays considerable intra-seasonal as well as inter-annual variability. Over the observational period, DOC concentrations were significantly negatively related to temperature and salinity, suggesting that outflow of Central Arctic waters carrying a high DOC load is the main control of DOC concentration in this region. POC concentration was not linked to temperature or salinity but tightly related to phytoplankton biomass as estimated from chlorophyll-a concentrations (Chl-a). For the years 2009–2017, no temporal trends in the depth-integrated (0–100 m) amounts of DOC and Chl-a were observed. In contrast, depth-integrated (0–100 m) amounts of POC, as well as the ratio [POC]:[TOC], decreased significantly over time. This suggests a higher partitioning of organic carbon into the dissolved phase. Potential causes and consequences of the observed changes in organic carbon stocks for food-web structure and CO2 sequestration are discussed.

Description: The Baltic Sea is prone to oxygen deficiency due to the restricted water exchange with the North Sea in coincidence with a high biological oxygen demand. The partitioning of organic carbon between respiration, accumulation and export is co-determined by phytoplankton primary production and its subsequent bacterial remineralization. Here, we investigated net phytoplankton primary production, heterotrophic bacterial biomass production and dark CO2 fixation by on-board incubations with radiolabeled tracers in the Baltic Proper and in the Gulf of Riga after the main spring bloom. Results show that low phytoplankton standing stocks of ≤1.6 μg chlorophyll a L–1 sustained net primary production of 161–724 mg C m–2 d–1 under nitrogen limitation. Estimates of bacterial carbon remineralization suggest that freshly produced organic carbon was supplied to the aphotic zone at all stations. In the southern Baltic Proper, net primary production exceeded the bacterial carbon demand in the surface mixed layer, suggesting that organic matter derived from nutrient-limited primary production was available for export to bacterial communities below the oxycline. On average, 46% of heterotrophic bacterial production was mediated in oxygen minimum zones, revealing the high importance of organic matter recycling under hypoxic and anoxic conditions for the carbon budget. Dark CO2 fixation of up to 4.33 μg C L–1 d–1 in sulfide-free waters equaled 9–54% of the co-inciding heterotrophic bacterial carbon demand and may have provided another organic carbon source for heterotrophic activity. Substantially higher dark CO2 fixation up to 25.46 μg C L–1 d–1 was determined in sulfidic waters. Since our study was conducted 5 months after the major Baltic inflow event in winter 2014/2015, potential effects of deep water ventilation could be investigated. In the Gotland Basin, heterotrophic bacterial production in renewed oxygen-rich bottom water was similar to that in the uplifted oxygen-deficient former bottom water, while it was significantly reduced in sulfidic waters. Hence, our results suggest that the removal of hydrogen sulfide by inflow events has a high potential to increase bacterial carbon remineralization.

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