Bioavailability of surface dissolved organic matter to aphotic bacterial communities in the Amundsen Sea Polynya, Antarctica

Antarctic seas, and particularly the Amundsen Sea Polynya, are some of the most productive oceanic regions on Earth. Ice-algal production during austral spring is followed by open-water pelagic production later in the season. Although ice-free growth accounts for a greater percentage of the annual net primary production, ice algae provide an important source of nutrients to organisms throughout the water column and benthos in areas and seasons when open-water production is insignificant. The objectives of this study were to assess the bioavailability of dissolved organic matter (DOM), sourced from ice algae or the chlorophyll maximum (chl max), to marine bacterioplankton and to determine the fate of carbon within these different DOM pools, including loss to respiration, incorporation into bacterial biomass and retention within the DOM pool itself. Nutrient concentrations and bacterial abundance, production, and cell volume were monitored during a 7-day bioassay study involving four treatments conducted shipboard in the Amundsen Sea Polynya, Antarctica. The greatest response in bacterial abundance and activity was observed when ice-algal meltwater was supplied to aphotic zone bacterioplankton collected from 170-m depth. However, bacterial growth efficiency was higher (24%) when chl max water was supplied to the same aphotic zone bacterial community compared to the bacterial growth efficiency of the ice-algal treatment (15%). Approximately 15% of dissolved organic carbon (DOC) from the ice-algal source and 18% from the chl max was consumed by aphotic bacterial communities over the relatively short, one-week incubation. In contrast, 65% of the dissolved organic nitrogen (DON) added as an integral part of the ice-algal DOM was consumed, but none of the DON supplied with chl max water was labile. This study underscores the importance of considering DOM sources when investigating or predicting changes in carbon and nitrogen cycling within the Amundsen Sea

A carbon budget for the Amundsen Sea Polynya, Antarctica: Estimating net community production and export in a highly productive polar ecosystem

Polynyas, or recurring areas of seasonally open water surrounded by sea ice, are foci for energy and material transfer between the atmosphere and the polar ocean. They are also climate sensitive, with both sea ice extent and glacial melt influencing their productivity. The Amundsen Sea Polynya (ASP) is the greenest polynya in the Southern Ocean, with summertime chlorophyll a concentrations exceeding 20 μg L−1. During the Amundsen Sea Polynya International Research Expedition (ASPIRE) in austral summer 2010–11, we aimed to determine the fate of this high algal productivity. We collected water column profiles for total dissolved inorganic carbon (DIC) and nutrients, particulate and dissolved organic matter, chlorophyll a, mesozooplankton, and microbial biomass to make a carbon budget for this ecosystem. We also measured primary and secondary production, community respiration rates, vertical particle flux and fecal pellet production and grazing. With observations arranged along a gradient of increasing integrated dissolved inorganic nitrogen drawdown (ΔDIN; 0.027–0.74 mol N m−2), changes in DIC in the upper water column (ranging from 0.2 to 4.7 mol C m−2) and gas exchange (0–1.7 mol C m−2) were combined to estimate early season net community production (sNCP; 0.2–5.9 mol C m−2) and then compared to organic matter inventories to estimate export. From a phytoplankton bloom dominated by Phaeocystis antarctica, a high fraction (up to ∼60%) of sNCP was exported to sub-euphotic depths. Microbial respiration remineralized much of this export in the mid waters. Comparisons to short-term (2–3 days) drifting traps and a year-long moored sediment trap capturing the downward flux confirmed that a relatively high fraction (3–6%) of the export from ∼100 m made it through the mid waters to depth. We discuss the climate-sensitive nature of these carbon fluxes, in light of the changing sea ice cover and melting ice sheets in the region

Interactive effects of elevated temperature and CO2 on nitrate, urea, and dissolved inorganic carbon uptake by a coastal California, USA, microbial community

Average global temperatures and carbon dioxide (CO2) levels are expected to increase in the coming decades. Implications for ocean ecosystems include shifts in microbial community structure and subsequent modifications to nutrient pathways. Studying how predicted future temperature and CO2 conditions will impact the biogeochemistry of the ocean is important because of the ocean’s role in regulating global climate. We determined how elevated temperature and CO2 affect uptake rates of nitrate, urea, and dissolved inorganic carbon (DIC) by 2 size classes (0.7-5.0 and \u3e5.0 µm) of a microbial assemblage collected from coastal California, USA. This microbial community was incubated for 10 d using an ecostat continuous culture system that supplied the microorganisms with either nitrate or urea as the dominant nitrogen source. Biomass parameters, nutrient concentrations, and uptake rates were measured throughout the experiment. In all treatments, urea uptake rates were greater than nitrate, and larger microorganisms had higher uptake rates than smaller microorganisms. Uptake rates of urea and DIC within both size fractions were higher at elevated temperature, and uptake rates of nitrate by smaller microorganisms increased with elevated CO2. These findings suggest that the rate at which nutrients cycle in temperate coastal waters will increase as temperature and CO2 levels rise and that the effect will vary between nitrogen substrates and different microorganisms

Stoichiometric N:P Ratios, Temperature, and Iron Impact Carbon and Nitrogen Uptake by Ross Sea Microbial Communities

The Southern Ocean is one of the most biologically important ecosystems on our planet. Microscopic plants, called phytoplankton, form the base of the food web in the Southern Ocean and play a direct role in regulating how much and how fast elements like nitrogen and carbon are cycled throughout the world ocean. The goal of this research was to determine how predicted changes in the environment will impact how fast phytoplankton use these elements. The conditions that we tested included elevated temperature, addition of iron, and the proportion of nitrogen to phosphorus in the seawater. These parameters were selected because temperatures are increasing in the Southern Ocean, and the relative availability of nutrients can alter what species of phytoplankton are present and how fast they grow. Phytoplankton were collected from two locations in the Ross Sea, Antarctica, and grown for a few weeks under experimental conditions. Our results demonstrate that all three parameters, warmer temperatures, the addition of iron, and changing nitrogen to phosphorus ratios will increase how fast phytoplankton use nitrogen and carbon, but the impact of elevated temperature and the addition of iron had a much larger impact than the nitrogen to phosphorus ratio

Seasonal nitrogen uptake and regeneration in the western coastal Arctic

Here, we present the first study to investigate the seasonal importance of amino acid-nitrogen (N) to Arctic near shore microbial communities. We measured primary productivity and the uptake of ammonium, nitrate, urea, and amino acids in two size fractions (\u3e 3 m and approximately 0.7-3 m), as well as ammonium regeneration and nitrification using N-15 and C-13 tracer approaches in the near-shore waters of the Chukchi Sea, during January, April, and August for two consecutive years. At discrete depths, nitrate comprised 46-78% of the total dissolved N pool during January and April but only 2-6% during August. Dissolved organic N (DON) concentrations increased between January and August though the carbon (C):N (mol:mol) of the DON pool declined. Of the substrates tested, amino acids supported the bulk of both N and C nutrition in both size fractions during January and April (ice-covered). Urea generally had the lowest uptake rate under ice-covered conditions; uptake of urea-C was only detectable in August. Though previous Arctic studies focused largely on nitrate, we found nitrate uptake was generally lower than other substrates tested. The sharp decline in nitrate concentration between April and August, however, indicates a drawdown of nitrate during that period. Rates of ammonium uptake were highest in August, when it was the dominant N substrate used. During all sample periods, rates of ammonium regeneration were sufficient to supply ammonium demand. Rates of nitrification varied between sample periods, however, with much higher rates seen in January and April

Preliminary estimate of contribution of Arctic nitrogen fixation to the global nitrogen budget

Dinitrogen (N-2) fixation is the source of all biologically available nitrogen on earth, and its presence or absence impacts net primary production and global biogeochemical cycles. Here, we report rates of 3.5-17.2 nmol N L-1 d(-1) in the ice-free coastal Alaskan Arctic to show that N-2 fixation in the Arctic Ocean may be an important source of nitrogen to a seasonally nitrogen-limited system. If widespread in surface waters over ice-free shelves throughout the Arctic, N-2 fixation could contribute up to 3.5 Tg N yr(-1) to the Arctic nitrogen budget. At these rates, N-2 fixation occurring in ice-free summer waters would offset up to 27.1% of the Arctic denitrification deficit and contribute an additional 2.7% to N-2 fixation globally, making it an important consideration in the current debate of whether nitrogen in the global ocean is in steady state. Additional investigations of high-latitude marine diazotrophic physiology are required to refine these N-2 fixation estimates

Symbiotic unicellular cyanobacteria fix nitrogen in the Arctic Ocean

Biological dinitrogen (N2) fixation is an important source of nitrogen (N) in low-latitude open oceans. The unusual N2-fixing unicellular cyanobacteria (UCYN-A)/haptophyte symbiosis has been found in an increasing number of unexpected environments, including northern waters of the Danish Straight and Bering and Chukchi Seas. We used nanoscale secondary ion mass spectrometry (nanoSIMS) to measure 15N2 uptake into UCYN-A/haptophyte symbiosis and found that UCYN-A strains identical to low-latitude strains are fixing N2 in the Bering and Chukchi Seas, at rates comparable to subtropical waters. These results show definitively that cyanobacterial N2 fixation is not constrained to subtropical waters, challenging paradigms and models of global N2 fixation. The Arctic is particularly sensitive to climate change, and N2 fixation may increase in Arctic waters under future climate scenarios

Microbial Community Response to Terrestrially Derived Dissolved Organic Matter in the Coastal Arctic

Warming at nearly twice the global rate, higher than average air temperatures are the new \u27normal\u27 for Arctic ecosystems. This rise in temperature has triggered hydrological and geochemical changes that increasingly release carbon-rich water into the coastal ocean via increased riverine discharge, coastal erosion, and the thawing of the semipermanent permafrost ubiquitous in the region. To determine the biogeochemical impacts of terrestrially derived dissolved organic matter (tDOM) on marine ecosystems we compared the nutrient stocks and bacterial communities present under ice-covered and ice-free conditions, assessed the lability of Arctic tDOM to coastal microbial communities from the Chukchi Sea, and identified bacterial taxa that respond to rapid increases in tDOM. Once thought to be predominantly refractory, we found that similar to 7% of dissolved organic carbon and similar to 38% of dissolved organic nitrogen from tDOM was bioavailable to receiving marine microbial communities on short 4 - 6 day time scales. The addition of tDOM shifted bacterial community structure toward more copiotrophic taxa and away from more oligotrophic taxa. Although no single order was found to respond universally (positively or negatively) to the tDOM addition, this study identified 20 indicator species as possible sentinels for increased tDOM. These data suggest the true ecological impact of tDOM will be widespread across many bacterial taxa and that shifts in coastal microbial community composition should be anticipated

Phytoplankton-bacterial interactions mediate micronutrient colimitation at the coastal Antarctic sea ice edge

Southern Ocean primary productivity plays a key role in global ocean biogeochemistry and climate. At the Southern Ocean sea ice edge in coastal McMurdo Sound, we observed simultaneous cobalamin and iron limitation of surface water phytoplankton communities in late Austral summer. Cobalamin is produced only by bacteria and archaea, suggesting phytoplankton-bacterial interactions must play a role in this limitation. To characterize these interactions and investigate the molecular basis of multiple nutrient limitation, we examined transitions in global gene expression over short time scales, induced by shifts in micronutrient availability. Diatoms, the dominant primary producers, exhibited transcriptional patterns indicative of co-occurring iron and cobalamin deprivation. The major contributor to cobalamin biosynthesis gene expression was a gammaproteobacterial population, Oceanospirillaceae ASP10-02a. This group also contributed significantly to metagenomic cobalamin biosynthesis gene abundance throughout Southern Ocean surface waters. Oceanospirillaceae ASP10-02a displayed elevated expression of organic matter acquisition and cell surface attachment-related genes, consistent with a mutualistic relationship in which they are dependent on phytoplankton growth to fuel cobalamin production. Separate bacterial groups, including Methylophaga, appeared to rely on phytoplankton for carbon and energy sources, but displayed gene expression patterns consistent with iron and cobalamin deprivation. This suggests they also compete with phytoplankton and are important cobalamin consumers. Expression patterns of siderophore-related genes offer evidence for bacterial influences on iron availability as well. The nature and degree of this episodic colimitation appear to be mediated by a series of phytoplankton-bacterial interactions in both positive and negative feedback loops