Microbial food web responses to light and nutrients beneath the coastal Arctic Ocean sea ice during the winter–spring transition

We measured the abundance and biomass of phototrophic and heterotrophic microbes in the upper mixed layer of the water column in ice-covered Franklin Bay, Beaufort Sea, Canada, from December 2003 to May 2004, and evaluated the influence of light and nutrients on these communities by way of a shipboard enrichment experiment. Bacterial cell concentrations showed no consistent trends throughout the sampling period, averaging (± SD) 2.4 (0.9) × 10⁸ cells L^− 1; integrated bacterial biomass for the upper mixed layer ranged from 1.33 mg C m^− 3 to 3.60 mg C m^− 3. Small cells numerically dominated the heterotrophic protist community in both winter and spring, but in terms of biomass, protists with a diameter > 10 µm generally dominated the standing stocks. Heterotrophic protist biomass integrated over the upper mixed layer ranged from 1.23 mg C m^− 3 to 6.56 mg C m^− 3. Phytoplankton biomass was low and variable, but persisted during the winter period. The standing stock of pigment-containing protists ranged from a minimum value of 0.38 mg C m^− 3 in winter to a maximal value of 6.09 mg C m^− 3 in spring and the most abundant taxa were Micromonas-like cells. These picoprasinophytes began to increase under the ice in February and their population size was positively correlated with surface irradiance. Despite the continuing presence of sea ice, phytoplankton biomass rose by more than an order of magnitude in the upper mixed layer by May. The shipboard experiment in April showed that this phototrophic increase in the community was not responsive to pulsed nutrient enrichment, with all treatments showing a strong growth response to improved irradiance conditions. Molecular (DGGE) and microscopic analyses indicated that most components of the eukaryotic community responded positively to the light treatment. These results show the persistence of a phototrophic inoculum throughout winter darkness, and the strong seasonal response by arctic microbial food webs to sub-ice irradiance in early spring.     

Number and phylogenetic affiliation of bacteria assimilating dimethylsulfoniopropionate and leucine in the ice-covered coastal Arctic Ocean

The ability of bacteria to assimilate sulfur from dimethylsulfoniopropionate (DMSP) was examined in the western Arctic Ocean by combining microautoradiography and fluorescence in situ hybridization (FISH). Assimilation of leucine was also measured for comparative purposes since leucine is considered a universal substrate for bacteria, which use it for protein synthesis. Samples were collected at 3 m depth, through a hole in the ice, in the CASES (Canadian Arctic Shelf Ecosystem Study) overwintering station in Franklin Bay (eastern Beaufort Sea) in March and May 2004 to compare two contrasting situations: winter and early spring. FISH counts indicated that the bacterial assemblage consisted of α- (up to 60% of the EUB positive cells), β- (up to 10%) and γ-proteobacteria (around 20%), and Bacteroidetes (up to 60%). The β-proteobacteria were not active with any of the two substrates tested. The remaining groups were much less efficient at assimilating DMSP-sulfur (5% of the cells) than leucine (20–35%) both in winter and in spring. Only the Roseobacter group of α-proteobacteria showed a similar assimilation of both substrates.     

Winter–spring dynamics in sea-ice carbon cycling in the coastal Arctic Ocean

An understanding of microbial interactions in first-year sea ice on Arctic shelves is essential for identifying potential responses of the Arctic Ocean carbon cycle to changing sea-ice conditions. This study assessed dissolved and particulate organic carbon (DOC, POC), exopolymeric substances (EPS), chlorophyll a, bacteria and protists, in a seasonal (24 February to 20 June 2004) investigation of first-year sea ice and associated surface waters on the Mackenzie Shelf. The dynamics of and relationships between different sea-ice carbon pools were investigated for the periods prior to, during and following the sea-ice-algal bloom, under high and low snow cover. A predominantly heterotrophic sea-ice community was observed prior to the ice-algal bloom under high snow cover only. However, the heterotrophic community persisted throughout the study with bacteria accounting for, on average, 44% of the non-diatom particulate carbon biomass overall the study period. There was an extensive accumulation of sea-ice organic carbon following the onset of the ice-algal bloom, with diatoms driving seasonal and spatial trends in particulate sea-ice biomass. DOC and EPS were also significant sea-ice carbon contributors such that sea-ice DOC concentrations were higher than, or equivalent to, sea-ice-algal carbon concentrations prior to and following the algal bloom, respectively. Sea-ice-algal carbon, DOC and EPS-carbon concentrations were significantly interrelated under high and low snow cover during the algal bloom (r values ≥ 0.74, p < 0.01). These relationships suggest that algae are primarily responsible for the large pools of DOC and EPS-carbon and that similar stressors and/or processes could be involved in regulating their release. This study demonstrates that DOC can play a major role in organic carbon cycling on Arctic shelves.     

Carbonate chemistry dynamics and carbon dioxide fluxes across the atmosphere–ice–water interfaces in the Arctic Ocean: Pacific sector of the Arctic

Climatic changes in the Northern Hemisphere have led to remarkable environmental changes in the Arctic Ocean, which is surrounded by permafrost. These changes include significant shrinking of sea-ice cover in summer, increased time between sea-ice break-up and freeze-up, and Arctic surface water freshening and warming associated with melting sea-ice, thawing permafrost, and increased runoff. These changes are commonly attributed to the greenhouse effect resulting from increased atmospheric carbon dioxide (CO₂) concentration and other non-CO₂ radiatively active gases (methane, nitrous oxide). The greenhouse effect should be most pronounced in the Arctic where the largest air CO₂ concentrations and winter–summer variations in the world for a clean background environment were detected. However, the air–land–shelf interaction in the Arctic has a substantial impact on the composition of the overlying atmosphere; as the permafrost thaws, a significant amount of old terrestrial carbon becomes available for biogeochemical cycling and oxidation to CO₂. The Arctic Ocean's role in determining regional CO₂ balance has been ignored, because of its small size (only  4% of the world ocean area) and because its continuous sea-ice cover is considered to impede gaseous exchange with the atmosphere so efficiently that no global climate models include CO₂ exchange over sea-ice. In this paper we show that: (1) the Arctic shelf seas (the Laptev and East-Siberian seas) may become a strong source of atmospheric CO₂ because of oxidation of bio-available eroded terrestrial carbon and river transport; (2) the Chukchi Sea shelf exhibits the strong uptake of atmospheric CO₂; (3) the sea-ice melt ponds and open brine channels form an important spring/summer air CO₂ sink that also must be included in any Arctic regional CO₂ budget. Both the direction and amount of CO₂ transfer between air and sea during open water season may be different from transfer during freezing and thawing, or during winter when CO₂ accumulates beneath Arctic sea-ice; (4) direct measurements beneath the sea ice gave two initial results. First, a drastic pCO₂ decrease from 410 μatm to 288 μatm, which was recorded in February–March beneath the fast ice near Barrow using the SAMI-CO₂ sensor, may reflect increased photosynthetic activity beneath sea-ice just after polar sunrise. Second, new measurements made in summer 2005 beneath the sea ice in the Central Basin show relatively high values of pCO₂ ranging between 425 μatm and 475 μatm, values, which are larger than the mean atmospheric value in the Arctic in summertime. The sources of those high values are supposed to be: high rates of bacterial respiration, import of the Upper Halocline Water (UHW) from the Chukchi Sea (CS) where values of pCO₂ range between 400 and 600 μatm, a contribution from the Lena river plume, or any combination of these sources.     

Methane release and coastal environment in the East Siberian Arctic shelf

In this paper we present 2 years of data obtained during the late summer period (September 2003 and September 2004) for the East Siberian Arctic shelf (ESAS). According to our data, the surface layer of shelf water was supersaturated up to 2500% relative to the present average atmospheric methane content of 1.85 ppm, pointing to the rivers as a strong source of dissolved methane which comes from watersheds which are underlain with permafrost. Anomalously high concentrations (up to 154 nM or 4400% supersaturation) of dissolved methane in the bottom layer of shelf water at a few sites suggest that the bottom layer is somehow affected by near-bottom sources. The net flux of methane from this area of the East Siberian Arctic shelf can reach up to 13.7 × 10⁴ g CH₄ km^− 2 from plume areas during the period of ice free water, and thus is in the upper range of the estimated global marine methane release. Ongoing environmental change might affect the methane marine cycle since significant changes in the thermal regime of bottom sediments within a few sites were registered. Correlation between calculated methane storage within the water column and both integrated salinity values (r = 0.61) and integrated values of dissolved inorganic carbon (DIC) (r = 0.62) suggest that higher concentrations of dissolved methane were mostly derived from the marine environment, likely due to in-situ production or release from decaying submarine gas hydrates deposits. The calculated late summer potential methane emissions tend to vary from year to year, reflecting most likely the effect of changing hydrological and meteorological conditions (temperature, wind) on the ESAS rather than riverine export of dissolved methane. We point out additional sources of methane in this region such as submarine taliks, ice complex retreat, submarine permafrost itself and decaying gas hydrates deposits.     

Planktonic community structure and carbon cycling in the Arabian Sea as a result of monsoonal forcing: the application of a generic model

The Arabian Sea exhibits a complex pattern of biogeochemical and ecological dynamics, which vary both seasonally and spatially. These dynamics have been studied using a one-dimensional vertical hydrodynamic model coupled to a complex ecosystem model, simulating the annual cycle at three contrasting stations. These stations are characterised by seasonally upwelling, mixed-layer-deepening and a-seasonal oligotrophic conditions, respectively, and coincide with extensively measured stations on the two JGOFS ARABESQUE cruises in 1994. The model reproduces many spatial and temporal trends in production, biomass, physical and chemical properties, both qualitatively and quantitatively and so gives insight into the main mechanisms responsible for the biogeochemical and ecological complexity. Monsoonal systems are typified by classical food web dynamics, whilst intermonsoonal and oligotrophic systems are dominated by the microbial loop. The ecosystem model (ERSEM), developed for temperate regions, is found to be applicable to the Arabian Sea system with little reparameterisation. Differences in in-situ physical forcing are sufficient to recreate contrasting eutrophic and oligotrophic systems, although the lack of lateral terms are probably the greatest source of error in the model. Physics, nutrients, light and grazing are all shown to play a role in controlling production and community structure. Small-celled phytoplanktons are predicted to be dominant and sub-surface chlorophyll maxima are robust centers of production during intermonsoon periods. Analysis of carbon fluxes indicate that physically driven outgassing of CO₂ predominates in monsoonal upwelling systems but ecological activity may significantly moderate CO₂ outgassing in the Arabian Sea interior.     

Ability of a “minimum” microbial food web model to reproduce response patterns observed in mesocosms manipulated with N and P, glucose, and Si

We compared an idealised mathematical model of the lower part of the pelagic food web to experimental data from a mesocosm experiment in which the supplies of mineral nutrients (nitrogen and phosphorous), bioavailable dissolved organic carbon (BDOC, as glucose), and silicate were manipulated. The central hypothesis of the experiment was that bacterial consumption of BDOC depends on whether the growth rate of heterotrophic bacteria is limited by organic-C or by mineral nutrients. In previous work, this hypothesis was examined qualitatively using a conceptual food web model. Here we explore the extent to which a “simplest possible” mathematical version of this conceptual model can reproduce the observed dynamics. The model combines algal–bacterial competition for mineral nutrients (phosphorous) and accounts for alternative limitation of bacterial and diatom growth rates by organic carbon and by silicate, respectively. Due to a slower succession in the diatom–copepod, compared to the flagellate–ciliate link, silicate availability increases the magnitude and extends the duration of phytoplankton blooms induced by mineral nutrient addition. As a result, Si interferes negatively with bacterial consumption of BDOC consumption by increasing and prolonging algal–bacterial competition for mineral nutrients. In order to reproduce the difference in primary production between Si and non-Si amended treatments, we had to assume a carbon overflow mechanism in diatom C-fixation. This model satisfactorily reproduced central features observed in the mesocosm experiment, including the dynamics of glucose consumption, algal, bacterial, and mesozooplankton biomass. While the parameter set chosen allows the model to reproduce the pattern seen in bacterial production, we were not able to find a single set of parameters that simultaneously reproduces both the level and the pattern observed for bacterial production. Profound changes in bacterial morphology and stoichiometry were reported in glucose-amended mesocosms. Our “simplest possible” model with one bacterial population with fixed stoichiometry cannot reproduce this, and we suggest that a more elaborate representation of the bacterial community is required for more accurate reproduction of bacterial production.