Terrestrial Inputs Shape Coastal Bacterial and Archaeal Communities in a High Arctic Fjord (Isfjorden, Svalbard)

The Arctic is experiencing dramatic changes including increases in precipitation, glacial melt, and permafrost thaw, resulting in increasing freshwater runoff to coastal waters. During the melt season, terrestrial runoff delivers carbon- and nutrient-rich freshwater to Arctic coastal waters, with unknown consequences for the microbial communities that play a key role in determining the cycling and fate of terrestrial matter at the land-ocean interface. To determine the impacts of runoff on coastal microbial (bacteria and archaea) communities, we investigated changes in pelagic microbial community structure between the early (June) and late (August) melt season in 2018 in the Isfjorden system (Svalbard). Amplicon sequences of the 16S rRNA gene were generated from water column, river and sediment samples collected in Isfjorden along fjord transects from shallow river estuaries and glacier fronts to the outer fjord. Community shifts were investigated in relation to environmental gradients, and compared to river and marine sediment microbial communities. We identified strong temporal and spatial reorganizations in the structure and composition of microbial communities during the summer months in relation to environmental conditions. Microbial diversity patterns highlighted a reorganization from rich communities in June toward more even and less rich communities in August. In June, waters enriched in dissolved organic carbon (DOC) provided a niche for copiotrophic taxa including Sulfitobacter and Octadecabacter. In August, lower DOC concentrations and Atlantic water inflow coincided with a shift toward more cosmopolitan taxa usually associated with summer stratified periods (e.g., SAR11 Clade Ia), and prevalent oligotrophic marine clades (OM60, SAR92). Higher riverine inputs of dissolved inorganic nutrients and suspended particulate matter also contributed to spatial reorganizations of communities in August. Sentinel taxa of this late summer fjord environment included taxa from the class Verrucomicrobiae (Roseibacillus, Luteolibacter), potentially indicative of a higher fraction of particle-attached bacteria. This study highlights the ecological relevance of terrestrial runoff for Arctic coastal microbial communities and how its impacts on biogeochemical conditions may make these communities susceptible to climate change

Light and freshwater discharge drive the biogeochemistry and microbial ecology in a sub-Arctic fjord over the Polar night

The polar night has recently received increased attention as a surprisingly active biological season. Yet, polar night microbial ecology is a vastly understudied field. To identify the physical and biogeochemical parameters driving microbial activity over the dark season, we studied a sub-Arctic fjord system in northern Norway from autumn to early spring with detailed monthly sampling. We focused on the impact of mixing, terrestrial organic matter input and light on microbial ecosystem dynamics. Our study highlights strong differences in the key drivers between spring, autumn, and winter. The spring bloom started in March in a fully mixed water column, opposing the traditional critical depth hypothesis. Incident solar radiation was the key driver maximum Chlorophyll was reached in April. The onset of the autumn phytoplankton bloom was controlled by vertical mixing, causing nutrient upwelling and dilution of zooplankton grazers, which had their highest biomass during this time. According to the dilution-recoupling hypothesis grazer dilution reduced grazing stress and allowed the fall bloom formation. Mixing at that time was initiated by strong winds and reduced stratification as a consequence of freezing temperatures and lower freshwater runoff. During the light-limited polar night, the primary production was extremely low but bacteria continued growing on decaying algae, their exudates and also allochthonous organic matter. A melting event in January could have increased input of organic matter from land, supporting a mid-winter bacterial bloom. In conclusion, polar night biogeochemistry and microbial ecology was not only driven by light availability, but strongly affected by variability in reshwater discharge and allochthonous carbon input. With climate change freshwater discharge will increase in the Arctic, which will likely increase importance of the dynamics described in this study

Light and freshwater discharge drive the biogeochemistry and microbial ecology in a sub-Arctic fjord over the Polar night

The polar night has recently received increased attention as a surprisingly active biological season. Yet, polar night microbial ecology is a vastly understudied field. To identify the physical and biogeochemical parameters driving microbial activity over the dark season, we studied a sub-Arctic fjord system in northern Norway from autumn to early spring with detailed monthly sampling. We focused on the impact of mixing, terrestrial organic matter input and light on microbial ecosystem dynamics. Our study highlights strong differences in the key drivers between spring, autumn, and winter. The spring bloom started in March in a fully mixed water column, opposing the traditional critical depth hypothesis. Incident solar radiation was the key driver maximum Chlorophyll was reached in April. The onset of the autumn phytoplankton bloom was controlled by vertical mixing, causing nutrient upwelling and dilution of zooplankton grazers, which had their highest biomass during this time. According to the dilution-recoupling hypothesis grazer dilution reduced grazing stress and allowed the fall bloom formation. Mixing at that time was initiated by strong winds and reduced stratification as a consequence of freezing temperatures and lower freshwater runoff. During the light-limited polar night, the primary production was extremely low but bacteria continued growing on decaying algae, their exudates and also allochthonous organic matter. A melting event in January could have increased input of organic matter from land, supporting a mid-winter bacterial bloom. In conclusion, polar night biogeochemistry and microbial ecology was not only driven by light availability, but strongly affected by variability in reshwater discharge and allochthonous carbon input. With climate change freshwater discharge will increase in the Arctic, which will likely increase importance of the dynamics described in this study

Early spring subglacial discharge plumes fuel under-ice primary production at a Svalbard tidewater glacier

Subglacial upwelling of nutrient-rich bottom water is known to sustain elevated summer primary production in tidewater-glacier-influenced fjord systems. However, the importance of subglacial upwelling during the early spring season has not been considered yet. We hypothesized that subglacial discharge under sea ice is present in early spring and that its flux is sufficient to increase phytoplankton primary productivity. We evaluated the effects of the submarine discharge on primary production in a seasonally fast-ice covered Svalbard fjord (Billefjorden) influenced by a tidewater outlet glacier in April and May 2019. We found clear evidence for subglacial discharge and upwelling. Although the estimated bottom-water entrainment factor (1.6) and total fluxes were lower than in summer studies, we still observed substantial impact on the fjord ecosystem and primary production at this time of the year. The subglacial discharge leads to a salinity-stratified surface water layer and sea ice formation with low bulk salinity and permeability. The combination of the stratified surface layer, a 2-fold higher under-ice irradiance due to thinner snow cover, and higher N and Si concentrations at the glacier front supported phytoplankton primary production 2 orders of magnitude higher (42.6 mg C m−2 d−1) compared to a marine reference site at the fast-ice edge. Reciprocal transplant experiments showed that nutrient supply increased phytoplankton primary production by approximately 30 %. The brackish-water sea ice at the glacier front with its low bulk salinity contained a reduced brine volume, limiting the inhabitable brine channel space and nutrient exchange with the underlying seawater compared to full marine sea ice. Microbial and algal communities were substantially different in subglacial-influenced water and sea ice compared to the marine reference site, sharing taxa with the subglacial outflow water. We suggest that with climate change, the retreat of tidewater glaciers in early spring could lead to decreased under-ice phytoplankton primary production. In contrast, sea ice algae production and biomass may become increasingly important, unless sea ice disappears first, in which case spring phytoplankton primary production may increase

Seasonal Variability in the Zooplankton Community Structure in a Sub-Arctic Fjord as Revealed by Morphological and Molecular Approaches

Phyto- and zooplankton in Arctic and sub-Arctic seas show very strong seasonal changes in diversity and biomass. Here we document the seasonal variability in the mesozooplankton community structure in a sub-Arctic fjord in Northern Norway based on monthly sampling between November 2018 and February 2020. We combined traditional morphological zooplankton identification with DNA metabarcoding of a 313 base pair fragment of the COI gene. This approach allowed us to provide the most detailed mesozooplankton species list known for this region across an entire year, including both holo- and meroplankton. The zooplankton community was dominated by small copepods throughout the sampling period both in terms of abundance and relative sequence counts. However, meroplankton was the most diverse group, especially within the phylum polychaeta. We identified four distinct periods based on the seasonal analysis of the zooplankton community composition. The pre-spring bloom period (February–March) was characterized by low abundance and biomass of zooplankton. The spring bloom (April) was characterized by the presence of Calanus young stages, cirripedia and krill eggs. The spring-summer period (May–August) was characterized by a succession of meroplankton and a relatively high abundance of copepods of the genus Calanus spp. Finally, the autumn-winter period (September–December) was characterized by a high copepod diversity and a peak in abundance of small copepods (e.g., Oithona similis, Acartia longiremis, Pseudocalanus acuspes, Pseudocalanus elongatus, Pseudocalanus moultoni, Pseudocalanus minutus). During this period, we also observed an influx of boreal warm-water species which were notably absent during the rest of the year. Both the traditional community analysis and metabarcoding were highly complementary and with a few exceptions showed similar trends in the seasonal changes of the zooplankton community structure

Modelling Silicate - Nitrate - Ammonium Co-Limitation of Algal Growth and the Importance of Bacterial Remineralisation Based on an Experimental Arctic Coastal Spring Bloom Culture Study

Arctic coastal ecosystems are rapidly changing due to climate warming, which makes modelling their productivity crucially important to better understand future changes. System primary production in these systems is highest during the pronounced spring bloom, typically dominated by diatoms. Eventually the spring blooms terminate due to silicon or nitrogen limitation. Bacteria can play an important role for extending bloom duration and total CO2 fixation through ammonium regeneration. Current ecosystem models often simplify the effects of nutrient co-limitations on algal physiology and cellular ratios and neglect bacterial driven regeneration, leading to an underestimation of primary production. Detailed biochemistry- and cell-based models can represent these dynamics but are difficult to tune in the environment. We performed a cultivation experiment that showed typical spring bloom dynamics, such as extended algal growth via bacteria ammonium remineralisation, and reduced algal growth and inhibited chlorophyll synthesis under silicate limitation, and gradually reduced nitrogen assimilation and chlorophyll synthesis under nitrogen limitation. We developed a simplified dynamic model to represent these processes. The model also highlights the importance of organic matter excretion, and post bloom ammonium accumulation. Overall, model complexity is comparable to other ecosystem models used in the Arctic while improving the representation of nutrient co-limitation related processes. Such model enhancements that now incorporate increased nutrient inputs and higher mineralization rates in a warmer climate will improve future predictions in this vulnerable system

Terrestrial Inputs Shape Coastal Bacterial and Archaeal Communities in a High Arctic Fjord (Isfjorden, Svalbard)

The Arctic is experiencing dramatic changes including increases in precipitation, glacial melt, and permafrost thaw, resulting in increasing freshwater runoff to coastal waters. During the melt season, terrestrial runoff delivers carbon- and nutrient-rich freshwater to Arctic coastal waters, with unknown consequences for the microbial communities that play a key role in determining the cycling and fate of terrestrial matter at the land-ocean interface. To determine the impacts of runoff on coastal microbial (bacteria and archaea) communities, we investigated changes in pelagic microbial community structure between the early (June) and late (August) melt season in 2018 in the Isfjorden system (Svalbard). Amplicon sequences of the 16S rRNA gene were generated from water column, river and sediment samples collected in Isfjorden along fjord transects from shallow river estuaries and glacier fronts to the outer fjord. Community shifts were investigated in relation to environmental gradients, and compared to river and marine sediment microbial communities. We identified strong temporal and spatial reorganizations in the structure and composition of microbial communities during the summer months in relation to environmental conditions. Microbial diversity patterns highlighted a reorganization from rich communities in June toward more even and less rich communities in August. In June, waters enriched in dissolved organic carbon (DOC) provided a niche for copiotrophic taxa including Sulfitobacter and Octadecabacter. In August, lower DOC concentrations and Atlantic water inflow coincided with a shift toward more cosmopolitan taxa usually associated with summer stratified periods (e.g., SAR11 Clade Ia), and prevalent oligotrophic marine clades (OM60, SAR92). Higher riverine inputs of dissolved inorganic nutrients and suspended particulate matter also contributed to spatial reorganizations of communities in August. Sentinel taxa of this late summer fjord environment included taxa from the class Verrucomicrobiae (Roseibacillus, Luteolibacter), potentially indicative of a higher fraction of particle-attached bacteria. This study highlights the ecological relevance of terrestrial runoff for Arctic coastal microbial communities and how its impacts on biogeochemical conditions may make these communities susceptible to climate change

Modeling silicate–nitrate–ammonium co-limitation of algal growth and the importance of bacterial remineralization based on an experimental Arctic coastal spring bloom culture study

Arctic coastal ecosystems are rapidly changing due to climate warming. This makes modeling their produc- tivity crucially important to better understand future changes. System primary production in these systems is highest dur- ing the pronounced spring bloom, typically dominated by di- atoms. Eventually the spring blooms terminate due to sili- con or nitrogen limitation. Bacteria can play an important role for extending bloom duration and total CO2 fixation through ammonium regeneration. Current ecosystem mod- els often simplify the effects of nutrient co-limitations on al- gal physiology and cellular ratios and simplify nutrient re- generation. These simplifications may lead to underestimations of primary production. Detailed biochemistry- and cell- based models can represent these dynamics but are difficult to tune in the environment. We performed a cultivation experiment that showed typical spring bloom dynamics, such as extended algal growth via bacterial ammonium remineralization, reduced algal growth and inhibited chlorophyll synthesis under silicate limitation, and gradually reduced nitrogen assimilation and chlorophyll synthesis under nitrogen limitation. We developed a simplified dynamic model to represent these processes. Overall, model complexity in terms of the number of parameters is comparable to the phytoplankton growth and nutrient biogeochemistry formulations in common ecosystem models used in the Arctic while improv- ing the representation of nutrient-co-limitation-related processes. Such model enhancements that now incorporate in- creased nutrient inputs and higher mineralization rates in a warmer climate will improve future predictions in this vulnerable system

Arctic marine fungi: biomass, functional genes, and putative ecological roles

Recent molecular evidence suggests a global distribution of marine fungi; however, the ecological relevance and corresponding biological contributions of fungi to marine ecosystems remains largely unknown. We assessed fungal biomass from the open Arctic Ocean by applying novel biomass conversion factors from cultured isolates to environmental sterol and CARD-FISH data. We found an average of 16.54 nmol m−3 of ergosterol in sea ice and seawater, which corresponds to 1.74 mg C m−3 (444.56 mg C m−2 in seawater). Using Chytridiomycota-specific probes, we observed free-living and particulate-attached cells that averaged 34.07 µg C m−3 in sea ice and seawater (11.66 mg C m−2 in seawater). Summed CARD-FISH and ergosterol values approximate 1.77 mg C m−3 in sea ice and seawater (456.23 mg C m−2 in seawater), which is similar to biomass estimates of other marine taxa generally considered integral to marine food webs and ecosystem processes. Using the GeoChip microarray, we detected evidence for fungal viruses within the Partitiviridae in sediment, as well as fungal genes involved in the degradation of biomass and the assimilation of nitrate. To bridge our observations of fungi on particulate and the detection of degradative genes, we germinated fungal conidia in zooplankton fecal pellets and germinated fungal conidia after 8 months incubation in sterile seawater. Ultimately, these data suggest that fungi could be as important in oceanic ecosystems as they are in freshwater environments