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River-ocean continuums are rich environments key to the transformations of organic matter and nutrients from many sources. Human impacts on these ecosystems can be local, upstream, or global. Particulate organic matter provides one tool for exploring these processes; inventories and carbon and nitrogen stable isotopes are useful indicators. This study conducted comprehensive field investigations from the upper reaches of the Yangtze River to the coastal waters to explore the distribution and spatial-temporal variation of particulate organic carbon (POC) and nitrogen (PON), and dissolved organic carbon (DOC) in surface waters from April–May (Spring) and October–November (Autumn) 2014. Carbon and nitrogen isotopes (δ13C and δ15 N) were used to assess organic matter sources. Numerous environmental factors were also measured. PON and POC exhibited similar spatial and temporal variations along the river mainstream. POC increased from the upper to lower reaches, similarly in both seasons; POC declined seaward from the river mouth. In contrast, PON showed greater seasonal and spatial variations along the continuum, with greater primary production in offshore areas. DOC along the continuum was higher in autumn, likely related to greater river discharge and warmer waters. The DOC/POC ratios by weight were >1 in most samples except for the ones collected from the turbidity maximum zone, suggesting that DOC contributed to a larger fraction of the total organic matter inventories. End-member mixing models revealed that the proportion of organic matter from upstream (allochthonous) sources was >70% in the mainstream during both seasons. In coastal waters, the proportion of allochthonous sources was only >57%. Compared with other major world rivers, DOC and allochthonous POC are especially high in the Yangtze River-ocean continuum. Human activities causing variations in terrestrial inputs are likely an important driving factor. In addition, the contribution of soil erosion caused by urbanization to riverine organic carbon cannot be ignored.

Numerous coastal polynyas fringe the Antarctic continent and strongly influence the productivity of Antarctic shelf systems. Of the 46 Antarctic coastal polynyas documented in a recent study, the Amundsen Sea Polynya (ASP) stands out as having the highest net primary production per unit area. Incubation experiments suggest that this productivity is partly controlled by the availability of dissolved iron (dFe). As a first step toward understanding the iron supply of the ASP, we introduce four plausible sources of dFe and simulate their steady spatial distribution using conservative numerical tracers. The modeled distributions replicate important features from observations including dFe maxima at the bottom of deep troughs and enhanced concentrations near the ice shelf fronts. A perturbation experiment with an idealized drawdown mimicking summertime biological uptake and subsequent resupply suggests that glacial meltwater and sediment-derived dFe are the main contributors to the pre-bloom dFe inventory in the top 100m of the ASP. The sediment-derived dFe depends strongly on the buoyancy-driven overturning circulation associated with the melting ice shelves (the ‘meltwater pump') to add dFe to the upper 300m of the water column. The results support the view that ice shelf melting plays an important direct and indirect role in the dFe supply and delivery to polynyas such as the ASP.

Each summer, large quantities of freshwater and associated dissolved and particulate material are released from the Greenland Ice Sheet (GrIS) into local fjords where they promote local phytoplankton growth. Whether the influx of freshwater and associated micronutrients in glacial meltwater is able to stimulate phytoplankton growth beyond the fjords is disputed, however. Here we show that the arrival of freshwater discharge from outlet glaciers from both southeast and southwest GrIS coincides with large-scale blooms in the Labrador Sea that extend over 300 km from the coast during summer. This summer bloom develops about a week after the arrival of glacial meltwater in early July and persists until the input of glacial meltwater slows in August or September, accounting for ~40% of annual net primary production for the area. In view of the absence of a significant change in the depth of the mixed layer associated with the arrival of glacial meltwater to the Labrador Sea, we suggest that the increase in phytoplankton biomass and productivity in summer is likely driven by a greater nutrient supply (most likely iron). Our results highlight that the ecological impact of meltwater from the GrIS likely extends far beyond the boundaries of the local fjords, encompassing much of the eastern Labrador Sea. Such impacts may increase if melting of the GrIS accelerates as predicted.

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 (> 3 μm and approximately 0.7–3 μm), as well as ammonium regeneration and nitrification using 15N and 13C 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.

The Arctic is warming at a rate nearly twice the global average, leading to thawing permafrost, increased coastal erosion, and enhanced delivery of riverine terrestrially-derived dissolved organic matter (tDOM) to coastal waters. This humic-rich tDOM has the ability to attenuate light required for photosynthesis and stimulate heterotrophic growth by supplying a source of labile organic carbon. Due to tDOM's high carbon to nitrogen (C : N) ratio, additional nitrogen is required for microorganisms to utilize this excess carbon for growth, thus exacerbating competition between autotrophs and heterotrophs for limiting nutrients and potentially reducing primary production. The effect of Arctic tDOM additions on nitrate uptake by two microplankton size fractions in the coastal Chukchi Sea was quantified using 15N tracer methods. To assess the biogeochemical vs. spectral impacts of tDOM, the uptake incubations were amended with either tDOM or light attenuating films that mimic light absorption by the tDOM. Nitrate uptake and primary production rates in the larger, predominantly phytoplankton, size fraction generally decreased with increasing tDOM additions. The change in light attenuation alone accounted for a ∼ 50% reduction in nitrate uptake. Responses in the smaller size fraction varied seasonally with tDOM additions stimulating uptake in spring and suppressing it in summer. The largest variation in summer nitrate uptake can be explained by the shared effect of biogeochemistry and light attenuation. Therefore, large increases in tDOM delivery currently occurring and predicted to increase in the coastal Arctic, could reduce primary production, broadly impact nitrogen and carbon cycling, and affect higher trophic levels.

As part of the ANACONDAS program, we investigated the role of the Amazon plume in stimulating offshore nitrogen fixation and export production during the river's high-discharge period (May–June 2010). Using the shipboard underway system, we performed high-resolution sampling of over 450,000 km2 of surface waters, characterizing the distribution of nutrients, phytoplankton, particulate organic matter (POM), and stable carbon and nitrogen isotopes of POM in the offshore plume. We found distinct regional variations in diazotroph communities, with the Diatom-Diazotroph Associations (DDA) Hemiaulus hauckii – Richelia intracellularis dominating the low N : P mesohaline waters to the northwest of the plume axis and Trichodesmium spp. primarily occupying oceanic waters to the east. Nutrient availability broadly shaped diazotroph distributions along the salinity gradient, but habitat longevity may also play a role in the finer-scale distributions of communities, particularly of DDAs. H. hauckii and Trichodesmium spp. affected the nitrogen and carbon budgets in fundamentally different ways within the plume-influenced regions, with H. hauckii making much greater contributions to the particulate nitrogen pool and to CO2 drawdown than Trichodesmium spp., leading to much higher export fluxes. Our findings provide an important constraint on the role of the Amazon plume in creating distinct niches and roles for diazotrophs in the nutrient and carbon budgets of the western tropical North Atlantic.

Marine dissolved organic matter (DOM) contains one of the largest exchangeable organic carbon pools on Earth. Riverine input represents an important source of DOM to the oceans, yet much remains to be learned about the fate of the DOM linking terrestrial to oceanic carbon cycles through rivers at the global scale. Here we use ultrahigh-resolution mass spectrometry to identify 184 molecular formulae that are indicators of riverine inputs (referred to as t-Peaks) and to track their distribution in the deep North Atlantic and North Pacific Oceans. The t-Peaks were found to be enriched in the Amazon River, to be highly correlated with known tracers of terrigenous input, and to be observed in all samples from four different rivers characterized by vastly different landscapes and vegetation coverage spanning equatorial (Amazon and Congo), subtropical (Altamaha), and Arctic (Kolyma) regions. Their distribution reveals that terrigenous organic matter is injected into the deep ocean by the global meridional overturning circulation, indicating that a fraction of the terrigenous DOM introduced by rivers contributes to the DOM pool observed in the deep ocean and to the storage of terrigenous organic carbon. This novel molecular approach can be used to further constrain the transfer of DOM from land to sea, especially considering that Fourier transform ion cyclotron resonance mass spectrometer analysis is becoming increasingly frequent in studies characterizing the molecular composition of DOM in lakes, rivers, and the ocean.

The Greenland ice sheet has undergone accelerating mass losses during recent decades. Freshwater runoff from ice melt can influence fjord circulation and dynamic1 and the delivery of bioavailable micronutrients to the ocean. It can also have climate implications, because stratification in the adjacent Labrador Sea may influence deep convection and the strength of the Atlantic meridional overturning circulation. Yet, the fate of the meltwater in the ocean remains unclear. Here, we use a high-resolution ocean model to show that only 1-15% of the surface meltwater runoff originating from southwest Greenland is transported westwards. In contrast, up to 50-60% of the meltwater runoff originating from southeast Greenland is transported westwards into the northern Labrador Sea, leading to significant salinity and stratification anomalies far from the coast. Doubling meltwater runoff, as predicted in future climate scenarios, results in a more-than-double increase in anomalies offshore that persists further into the winter. Interannual variability in offshore export of meltwater is tightly related to variability in wind forcing. The new insight that meltwaters originating from the west and east coasts have different fates indicates that future changes in mass loss rates and surface runoff will probably impact the ocean differently, depending on their Greenland origins.

The Amundsen Sea Polynya (ASP) is distinguished by having the highest net primary production per unit area in the coastal Antarctic. Recent studies have related this high productivity to the presence of fast-melting ice shelves, but the mechanisms involved are not well understood. In this study we describe the first numerical model of the ASP to represent explicitly the ocean-ice interactions, nitrogen and iron cycles, and the coastal circulation at high resolution. The study focuses on the seasonal cycle of iron and carbon, and the results are broadly consistent with field observations collected during the summer of 2010-2011. The simulated biogeochemical cycle is strongly controlled by light availability(dictated by sea ice, phytoplankton self-shading, and variable sunlight). The micronutrient iron exhibits strong seasonality, where scavenging by biogenic particles and remineralization play large compensating roles. Lateral fluxes of iron are also important to the iron budget, and our results confirm the key role played by inputs of dissolved iron from the buoyancy-driven circulation of melting ice shelf cavities (the "meltwater pump"). The model suggests that westward flowing coastal circulation plays two important roles: it provides additional iron to the ASP and it collects particulate organic matter generated by the bloom and transports it to the west of the ASP. As a result, maps of vertical particulate organic matter fluxes show highest fluxes in shelf regions located west of the productive central ASP. Overall, these model results improve our mechanistic understanding of the ASP bloom, while suggesting testable hypotheses for future field efforts.

Riverine nutrients acting in concert with local hydrographic conditions create distinct ecological niches for phytoplankton communities across river-ocean continuums. Here we compare two of the world’s largest river-ocean systems, the Amazon River Plume (ARP) which outflows into the Western Tropical North Atlantic (WTNA) and the Changjiang Diluted Water (CDW) which empties into the East China Sea to show how distinctly different N: P ratios of their source waters, shape phytoplankton communities along the river-ocean continuum. Sampling in the relatively unpolluted surface waters of the ARP during peak river discharge revealed that phytoplankton communities along the river-ocean continuum were strongly controlled by Dissolved Inorganic Nitrogen (DIN, nitrate plus nitrite) concentrations which were low or beyond detectable, while Dissolved Inorganic Phosphorous (DIP, phosphate) was not limiting. Downstream of the plume, nutrient concentrations at the surface were far below the Redfield N:P ratio of 16 : 1, and in this severely nitrogen limited but silica replete mesohaline region, Diatom-Diazotroph Associations (DDA) such as Hemiaulus hauckii-Richelia proliferated, while offshore in nitrate poor oceanic waters, Trichodesmium spp. and Prochlorococcus thrived. In contrast, in the CDW, a system which has witnessed a substantial rise in anthropogenic nitrogen inputs from human pressures within the Changjiang River system, phytoplankton community structure was regulated primarily by DIP which was rapidly consumed in the inner shelf. Data from a series of summer-time cruises in the mid-shelf and offshore regions of the East China Sea, at peak discharge, established that when DIP limitation was ameliorated by on-shelf upwelling of DIP rich Kuroshio Intermediate Waters, diatoms and dinoflagellates dominated. Conversely, during years of heavy discharge, the westward flowing CDW plume was severely DIP limited, probably because water column stratification dampened upwelling of subsurface waters. The consequent phosphate limitation led to the proliferation of small phytoplankton such as Chlorophytes and Cyanobacteria. The absence of diazotrophs in the CDW, leads us to hypothesize that river-ocean continuums, whose source waters are heavily impacted by anthropogenic activities and with nutrient concentrations substantially in excess of Redfield ratios, may not support diatoms on account of DIP limitation nor DDAs and diazotrophs because of excess DIN.

Coastal oceans link terrestrial and marine carbon cycles. Yet, carbon sources and sinks in these biomes remain poorly understood. Here, we explore the dynamics of dissolved organic matter (DOM) along the Amazon River-to-ocean continuum from the lower mainstem at Obidos to the open ocean of the western tropical North Atlantic. We molecularly characterized DOM via ultrahigh-resolution Fourier-transform ion cyclotron resonance mass spectrometry (FT-ICR-MS), determined DOM stable carbon isotopes, and interpreted the data in the context of bacterial abundance and production, phytoplankton biomass and composition. Multivariate analysis revealed that the DOM molecular variability in the plume was mainly influenced by the input of terrigenous DOM. Incubation experiments with water from close to the river mouth showed that photo- and bio-degradation preferentially removed 13C-depleted and 13C-enriched terrigenous DOM, respectively. However, there was no significant quantitative change in the total amount of dissolved organic carbon (DOC) over five days. This result suggests that most of the reactive DOM had already been bio-degraded upstream within the river and that photo-degradation was diminished in the turbid plume close to the river mouth. Terrigenous DOM therefore appeared to be relatively non-reactive nearshore. In the less turbid offshore plume, enhanced light penetration stimulated growth of phytoplankton and increased bacterial production. Although marine DOM compounds became relatively enriched, bulk DOC concentrations were 9 to 30% below levels expected from conservative mixing of river and ocean endmembers suggesting that quantitative removal of terrigenous DOM was not compensated by marine DOM production. We propose that removal of terrigenous DOM in the outer plume may be enhanced by (i) bio-degradation primed by reactive algal DOM, (ii) photo-degradation, which may further break down DOM into more bio-available forms, and possibly (iii) sorption of DOM to sinking particles.

River systems play a pivotal role in transporting and transforming organic carbon (OC) fixed by terrestrial primary production. However, there is a fundamental gap in our understanding of the connectivity of terrestrial, aquatic, and marine carbon budgets due to a lack of measurements along the lower (i.e. tidally-influenced) reaches of large river systems. For example, all estimates of carbon fluxes from the world's largest river, the Amazon, are based on measurements made at and upstream of Obidos, roughly 900 km from the mouth. Here we examine the evolution of OC concentrations and composition from Obidos to two discreet channels near the mouth of the Amazon River during five cruises from 2010 to 2012. OC characteristics of the Tapajos River, which enters the Amazon River downstream of Obidos, and the Tocantins River, which mixes with the Amazon River plume in the Atlantic Ocean, were also assessed. The average concentration of particulate organic carbon (POC) across the two main channels near the mouth was 0.6 ± 0.3 mg L − 1 during the study period, decreasing from 1.2 ± 1.0 mg L − 1 at Obidos. Average dissolved organic carbon (DOC) concentrations, on the other hand, increased from 3.9 ± 0.6 mg L − 1 at Obidos to 4.2 ± 0.9 mg L − 1 across the mouth. The discharge of total OC to the ocean was composed of 89 ± 3% dissolved load, compared to 76 ± 13% at Obidos. Measurements of bulk OC stable isotopic signatures, chlorophyll a concentrations, and lignin phenol abundance were used to assess unique OC sources along the continuum. For example, the stable isotopic composition of POC and DOC indicated a replacement of highland forest-derived OC with lowland and floodplain-derived OC from Obidos to the mouth. Likewise, lignin phenol signatures showed an increase in the degradation state of vascular plant-derived OC from Obidos to the mouth. Results from this study illustrate that the abundance and composition of OC continue to evolve along the lower reaches of large tropical rivers, which has significant implications on estimations of geochemical fluxes to the ocean.

Constraining the fate of dissolved organic matter (DOM) delivered by rivers is a key to understand the global carbon cycle, since DOM mineralization directly influences air-sea CO2 exchange and multiple biogeochemical processes. The Amazon River exports large amounts of DOM, and yet the fate of this material in the ocean remains unclear. Here we investigate the molecular composition and transformations of DOM in the Amazon River-ocean continuum using ultrahigh resolution mass spectrometry and geochemical and biological tracers. We show that there is a strong gradient in source and composition of DOM along the continuum, and that dilution of riverine DOM in the ocean is the dominant pattern of variability in the system. Alterations in DOM composition are observed in the plume associated with the addition of new organic compounds by phytoplankton and with bacterial and photochemical transformations. The relative importance of each of these drivers varies spatially and is modulated by seasonal variations in river discharge and ocean circulation. We further show that a large fraction (50–76%) of the Amazon River DOM is surprisingly stable in the coastal ocean. This results in a globally significant river plume with a strong terrigenous signature and in substantial export of terrestrially derived organic carbon from the continental margin, where it can be entrained in the large-scale circulation and potentially contribute to the long-term storage of terrigenous production and to the recalcitrant carbon pool found in the deep ocean.

Analysis of metatranscriptomic and metagenomic datasets from the lower reaches of the Amazon River between Óbidos and the river mouth revealed microbial transcript and gene pools dominated by Actinobacteria, Thaumarchaeota, Bacteroidetes, Acidobacteria, Betaproteobacteria, and Planctomycetes. Three mainstem stations spanning a 625 km reach had similar gene expression patterns (transcripts gene copy−1) across a diverse suite of element cycling genes, but two tributary-influenced stations at the mouth of the Tapajós River and near the Tocantins River at Belém had distinct transcriptome composition and expression ratios, particularly for genes encoding light-related energy capture (higher) and iron acquisition and ammonia oxidation (lower). Environmental parameters that were useful predictors of gene expression ratios included concentrations of lignin phenols, suspended sediments, nitrate, phosphate, and particulate organic carbon and nitrogen. Similar to the gene expression data, these chemical properties reflected highly homogeneous mainstem stations punctuated by distinct tributary-influenced stations at Tapajós and Belém. Although heterotrophic processes were expected to dominate in the lower Amazon, transcripts from photosynthetic bacteria were abundant in tributary-influenced regions, and transcripts from Thaumarcheota taxa genetically capable of chemosynthetic ammonia oxidation accounted for up to 21% of the transcriptome at others. Based on regressions of transcript numbers against gene numbers, expression ratios of Thaumarchaeota populations were largely unchanged within the mainstem, suggesting a relatively minor role for gene regulation. These quantitative gene and transcript inventories detail a diverse array of energy acquisition strategies and metabolic capabilities for bacteria and archaea populations of the world's largest river system.

Biogeochemical rate processes in the Arctic are not currently well constrained, and there is very limited information on how rates may change as the region warms. Here we present data on the sensitivity of ammonium (NH4+) uptake and nitrification rates to short-term warming. Samples were collected from the Chukchi Sea off the coast of Barrow, Alaska, during winter, spring, and summer and incubated for 24 h in the dark with additions of 15NH4+ at −1.5, 6, 13, and 20°C. Rates of NH4+ uptake and nitrification were measured in conjunction with bacterial production. In all seasons, NH4+ uptake rates were highest at temperatures similar to current summertime conditions but dropped off with increased warming, indicative of psychrophilic (i.e., cold-loving) microbial communities. In contrast, nitrification rates were less sensitive to temperature and were higher in winter and spring compared to summer. These findings suggest that as the Arctic coastal ecosystem continues to warm, NH4+ assimilation may become increasingly important, relative to nitrification, although the magnitude of NH4+ assimilation would be still be lower than nitrification.

The Amazon River Plume delivers freshwater and nutrients to an otherwise oligotrophic western tropical North Atlantic (WTNA) Ocean. Plume waters create conditions favorable for carbon and nitrogen fixation, and blooms of diatoms and their diazotrophic cyanobacterial symbionts have been credited with significant CO 2 uptake from the atmosphere. The fate of the carbon, however, has been measured previously by just a few moored or drifting sediment traps, allowing only speculation about the full extent of the plume's impact on carbon flux to the deep sea. Here, we used surface (0.5 m) sediment cores collected throughout the Demerara Slope and Abyssal Plain, at depths ranging from 1800 to 5000 m, to document benthic diagenetic processes indicative of carbon flux. Pore waters were extracted from sediments using both mm- and cm-scale extraction techniques. Profiles of nitrate (NO 3 ) and silicate (Si(OH) 4 ) were modeled with a diffusion-reaction equation to determine particulate organic carbon (POC) degradation and biogenic silica (bSi) remineralization rates. Model output was used to determine the spatial patterns of POC and bSi arrival at the sea floor. Our estimates of POC and Si remineralization fluxes ranged from 0.16 to 1.92 and 0.14 to 1.35 mmol m −2 d −1 , respectively. A distinct axis of POC and bSi deposition on the deep sea floor aligned with the NW axis of the plume during peak springtime flood. POC flux showed a gradient along this axis with highest fluxes closest to the river mouth. bSi had a more diffuse zone of deposition and remineralization. The impact of the Amazon plume on benthic fluxes can be detected northward to 10°N and eastward to 47°W, indicating a footprint of nearly 1 million km 2 . We estimate that 0.15 Tmol C y −1 is remineralized in abyssal sediments underlying waters influenced by the Amazon River. This constitutes a relatively high fraction (~7%) of the estimated C export from the region.; the plume thus has a demonstrable impact on C org export in the western Atlantic. Benthic fluxes under the plume were comparable to and in some cases greater than those observed in the eastern equatorial Atlantic, the southeastern Atlantic, and the Southern Ocean.

18 pages, 6 figures, 1 table, supporting information https://doi.org/10.1002/ecs2.2641, During austral spring and summer, the coastal Antarctic experiences a sharp increase in primary production and a steepening of biotic and abiotic gradients that result from increased solar radiation and retreating sea ice. In one of the largest seasonally ice-free regions, the Amundsen Sea Polynya, pelagic samples were collected from 15 sites during a massive Phaeocystis antarctica bloom in 2010/2011. Along with a suite of other biotic and abiotic measurements, bacterioplankton were collected and analyzed for community structure by pyrosequencing of the 16S rRNA gene. The aims were to identify patterns in diversity and composition of heterotrophic bacterioplankton and to test mechanistic hypotheses for explaining these differences along variations in depth, water mass, phytoplankton biomass, and organic and inorganic nutrients. The overall goal was to clarify the relationship between primary producers and bacterioplankton community structure in the Southern Ocean. Results suggested that both epipelagic and mesopelagic bacterioplankton communities were structured by phytoplankton blooming in the euphotic zone. As chlorophyll a (chl-a) increased in surface waters, the abundance of surface bacterioplankton increased, but their diversity decreased. Similarity in bacterioplankton community composition between surface-water sites increased as the bloom progressed, suggesting that algal blooms may homogenize surface-water bacterioplankton communities at larger spatial scales. Below the euphotic zone, the opposite relationship was found. Mesopelagic bacterioplankton diversity increased with increasing chl-a in the overlying surface waters. This shift may be promoted by several factors including local increase in organic and inorganic nutrients from particles sinking out of the euphotic zone, an increase in niche differentiation associated with the particle flux, interactions with deep-dwelling macrozooplankton, and release from competition with primary producers. Additional multivariate analyses of bacterioplankton community structure and nutrient concentrations revealed distinct depth horizons, with bacterioplankton communities having maximum alpha and beta diversity just below the euphotic zone, while nutrient composition gradually homogenized with increasing depth. Our results provide evidence for bloom-driven (bottom-up) control of bacterioplankton community diversity in the coastal Southern Ocean and suggest mechanisms whereby surface processes can shape the diversity of bacterioplankton communities at great depth, The study was funded by the Swedish Research Council (grants to SB and LR) and the U.S. National Science Foundation through the ASPIRE project (ANT‐0839069)

An Advanced Laser Fluorometer (ALF) capable of discriminating several phytoplankton pigment types was utilized in conjunction with microscopic data to map the distribution of phytoplankton communities in the Amazon River plume in May–June-2010, when discharge from the river was at its peak. Cluster analysis and Non-metric Multi-Dimensional Scaling (NMDS) helped distinguish three distinct biological communities that separated largely on the basis of salinity gradients across the plume. These three communities included an ‘‘estuarine type’’ comprised of a high biomass mixed population of diatoms, cryptophytes and green-water Synechococcus spp. located upstream of the plume, a ‘‘mesohaline type’’ made up largely of communities of Diatom-Diazotroph Associations (DDAs) and located in the northwestern region of the plume and an ‘‘oceanic type’’ in the oligotrophic waters outside of the plume made up of Trichodesmium and Synechococcus spp. Although salinity appeared to have a substantial influence on the distribution of different phytoplankton groups, ALF and microscopic measurements examined in the context of the hydro-chemical environment of the river plume, helped establish that the phytoplankton community structure and distribution were strongly controlled by inorganic nitrate plus nitrite (NO3 +N O 2) availability whose concentrations were low throughout the plume. Towards the southern, low-salinity region of the plume, NO3 +N O 2 supplied by the onshore flow of subsurface (80 m depth) water, ensured the continuous sustenance of the mixed phytoplankton bloom. The large drawdown of SiO3 and PO4 associated with this ‘‘estuarine type’’ mixed bloom at a magnitude comparable to that observed for DDAs in the mesohaline waters, leads us to contend that, diatoms, cryptophytes and Synechococcus spp., fueled by the offshore influx of nutrients also play an important role in the cycling of nutrients in the Amazon River plume.

[1] The Amazon River Plume spreads across the tropical North Atlantic creating a barrier to vertical mixing. Here using a 1/6° HYCOM model and data from three research cruises in May–June 2010, September–October 2011, and July 2012, we investigate the pathways and properties of the plume. Four plume pathways for export of freshwater from the western tropical North Atlantic are identified. These consist of direct and indirect pathways to the northwest, and eastward pathways toward the subtropical gyre and toward Africa in the North Equatorial Counter Current. Because of the seasonality and cooccurrence of these pathways, plume characteristics are highly variable. Two pathways export water to the Caribbean, however the time scales associated with those direct and indirect pathways (3 versus 6+ months) differ, leading to different salinity characteristics of the plume water. Models results show that the Amazon river and tropical precipitation have similar magnitude impact on the observed seasonal cycle of freshwater within the western tropical Atlantic and at the 8°N, 38°W PIRATA mooring. Freshwater associated with the Amazon also influences surface salinity in winter as far as 20W in the model. The mean plume salinity minimum leads maximum discharge, highlighting the importance of currents and advection rather than discharge in maintaining plume properties. Plume pathways are tied to the underlying current structure, with the North Equatorial Counter Current jet preventing direct freshwater transport into the southern hemisphere. The plume influences underlying currents as well, generating vertical current shear that leads to enhanced eddy stirring and mixing in the model simulations.

Heterotrophic bacteria play a key role in marine carbon cycling, and understanding their activities in polar systems is important for considering climate change impacts there. One goal of the ASPIRE project was to examine the relationship between the phytoplankton bloom and bacterial heterotrophy in the Amundsen Sea Polynya (ASP). Bacterial abundance, production (BP), respiration, growth efficiency, and extracellular enzyme activity (EEA) were compared to nutrient and organic matter inventories, chlorophyll a (Chl a), viral and microzooplankton abundance, and net primary production (NPP). Bacterial production and respiration clearly responded (0.04–4.0 and 10–53 µg C L−1 d−1, respectively) to the buildup of a massive Phaeocystis antarctica bloom (Chl a: 0.2–22 µg L−1), with highest rates observed in the central polynya where Chl a and particulate organic carbon (POC) were greatest. The highest BP rates exceeded those reported for the Ross Sea or any other Antarctic coastal system, yet the BP:NPP ratio (2.1–9.4%) was relatively low. Bacterial respiration was also high, and growth efficiency (2–27%; median = 10%) was similar to oligotrophic systems. Thus, the integrated bacterial carbon demand (0.8–2.8 g C m−2 d−1) was a high fraction (25–128%; median = 43%) of NPP during bloom development. During peak bloom, activity was particle-associated: BP and EEA correlated well with POC, and size fractionation experiments showed that the larger size fraction (> 3 µm) accounted for a majority (∼ 75%) of the BP. The community was psychrophilic, with a 5x reduction in BP when warmed to 20°C. In deeper waters, respiration remained relatively high, likely fueled by the significant downward particle flux in the region. A highly active, particle-associated, heterotrophic microbial community clearly responded to the extraordinary phytoplankton bloom in the ASP, likely limiting biological pump efficiency during the early season.

6 pages, 4 figures, The Antarctic and Arctic regions offer a unique opportunity to test factors shaping biogeography of marine microbial communities because these regions are geographically far apart, yet share similar selection pressures. Here, we report a comprehensive comparison of bacterioplankton diversity between polar oceans, using standardized methods for pyrosequencing the V6 region of the small subunit ribosomal (SSU) rRNA gene. Bacterial communities from lower latitude oceans were included, providing a global perspective. A clear difference between Southern and Arctic Ocean surface communities was evident, with 78% of operational taxonomic units (OTUs) unique to the Southern Ocean and 70% unique to the Arctic Ocean. Although polar ocean bacterial communities were more similar to each other than to lower latitude pelagic communities, analyses of depths, seasons, and coastal vs. open waters, the Southern and Arctic Ocean bacterioplankton communities consistently clustered separately from each other. Coastal surface Southern and Arctic Ocean communities were more dissimilar from their respective open ocean communities. In contrast, deep ocean communities differed less between poles and lower latitude deep waters and displayed different diversity patterns compared with the surface. In addition, estimated diversity (Chao1) for surface and deep communities did not correlate significantly with latitude or temperature. Our results suggest differences in environmental conditions at the poles and different selection mechanisms controlling surface and deep ocean community structure and diversity. Surface bacterioplankton may be subjected to more short-term, variable conditions, whereas deep communities appear to be structured by longer water-mass residence time and connectivity through ocean circulation, We thank the members of field teams, shipboard crews, and logistics support personnel from all national polar programs involved in sample collection, without whom this study would not have been possible. The Census of Antarctic Marine Life, funded by the Sloan Foundation, facilitated this collaboration. Pyrosequencing was provided by the International Census of Marine Microbes (ICoMM) with financial support from a W. M. Keck Foundation award to the Marine Biological Laboratory in Woods Hole. Funding to support sample collection was provided by the Institut Français pour la Recherche et la Technologie Polaires (J.-F.G.); the Spanish Ministry of Education and Science (C.P.-A.); the New Zealand International Polar Year-Census of Antarctic Marine Life Project [Phases 1 (So001IPY) and 2 (IPY2007-01); to E.W.M.); the Natural Sciences and Engineering Council (NSERC) of Canada (C.L.); National Science Foundation Grants OPP-0124733 (to D.L.K.), ANT-0632389 (to A.E.M.), and ANT-0741409 (to P.L.Y.); and the Swedish Polar Research Secretariat (S.B.)

The South Atlantic Bight (SAB) has been a focus for the study of continental shelf ecosystem respiration during the past two decades. However, two questions concerning respiration in this area have yet to be answered. First, why do previous estimates of respiration in the SAB exceed measured carbon fixation rates by almost an order of magnitude? Second, considering that bacteria are responsible for most of the pelagic community respiration in the SAB, why is respiration almost uniform from the coastline to the shelf break, while bacterial production estimates decrease offshore? This study addresses these critical questions by presenting new pelagic community respiration data that were collected across the entire width of the continental shelf off Georgia, USA from June 2003 to May 2006. The respiration was calculated as in vitro changes of dissolved oxygen and dissolved inorganic carbon concentrations during deck incubations. The measured respiration rates ranged from 0.3(±0.1) to 21.2(±1.4) mmol m−3 day−1. They followed a clear seasonal pattern, being lowest over the entire shelf in winter and reaching maxima in summer. Summertime respiration rates were highest on the inner shelf and decreased with distance offshore. Consistent with this trend, bacterial abundance measurements taken during the sampling month of July 2005 followed a pattern of seaward decline. The SAB organic carbon fluxes calculated from the respiration data are close to the estimates for primary production, which resolves a long-standing mystery regarding perceived carbon imbalance in the SAB.

The fresh water discharged by large rivers such as the Amazon is transported hundreds to thousands of kilometers away from the coast by surface plumes. The nutrients delivered by these river plumes contribute to enhanced primary production in the ocean, and the sinking flux of this new production results in carbon sequestration. Here, we report that the Amazon River plume supports N 2 fixation far from the mouth and provides important pathways for sequestration of atmospheric CO 2 in the western tropical North Atlantic (WTNA). We calculate that the sinking of carbon fixed by diazotrophs in the plume sequesters 1.7 Tmol of C annually, in addition to the sequestration of 0.6 Tmol of C yr −1 of the new production supported by NO 3 delivered by the river. These processes revise our current understanding that the tropical North Atlantic is a source of 2.5 Tmol of C to the atmosphere [Mikaloff-Fletcher SE, et al. (2007) Inverse estimates of the oceanic sources and sinks of natural CO 2 and the implied oceanic carbon transport. Global Biogeochem Cycles 21, doi:10.1029/2006GB002751]. The enhancement of N 2 fixation and consequent C sequestration by tropical rivers appears to be a global phenomenon that is likely to be influenced by anthropogenic activity and climate change.

Yager, Patricia L. ... et al.-- Special feature: ASPIRE: The Amundsen Sea Polynya International Research Expedition.-- 36 pages, 11 figures, 9 tables, supplementary data http://www.bco-dmo.org/project/2132, 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, This project was funded by NSF Office of Polar Programs, Antarctic Organisms and Ecosystems (ANT-0839069 to PY, ANT-0838995 to RMS, ANT-0838975 to SS, ANT-0839012 to HWD, and ANT-0944727 to K Arrigo for support of ACA, GLvD and KEL). We also acknowledge support for participation of AFP by NSF (ANT-08-39012 to HWD and ANT 15-42220 to AFP) and for Swedish collaborators from the Swedish Research Council (#2008-6430 to S. Bertilsson and L. Riemann, and #2008-6429 to P.-O. Moksness and J. Havenhand, which supported participation of RS and SK)

A novel Amazonian reef biome was discovered, encompassing large rhodolith and sponge beds under low light, low oxygen, and high POC., Large rivers create major gaps in reef distribution along tropical shelves. The Amazon River represents 20% of the global riverine discharge to the ocean, generating up to a 1.3 × 106–km2 plume, and extensive muddy bottoms in the equatorial margin of South America. As a result, a wide area of the tropical North Atlantic is heavily affected in terms of salinity, pH, light penetration, and sedimentation. Such unfavorable conditions were thought to imprint a major gap in Western Atlantic reefs. We present an extensive carbonate system off the Amazon mouth, underneath the river plume. Significant carbonate sedimentation occurred during lowstand sea level, and still occurs in the outer shelf, resulting in complex hard-bottom topography. A permanent near-bottom wedge of ocean water, together with the seasonal nature of the plume’s eastward retroflection, conditions the existence of this extensive (~9500 km2) hard-bottom mosaic. The Amazon reefs transition from accretive to erosional structures and encompass extensive rhodolith beds. Carbonate structures function as a connectivity corridor for wide depth–ranging reef-associated species, being heavily colonized by large sponges and other structure-forming filter feeders that dwell under low light and high levels of particulates. The oxycline between the plume and subplume is associated with chemoautotrophic and anaerobic microbial metabolisms. The system described here provides several insights about the responses of tropical reefs to suboptimal and marginal reef-building conditions, which are accelerating worldwide due to global changes.

The Amundsen Sea Polynya is characterized by large phytoplankton blooms, which makes this region disproportionately important relative to its size for the biogeochemistry of the Southern Ocean. In situ data on phytoplankton are limited, which is problematic given recent reports of sustained change in the Amundsen Sea. During two field expeditions to the Amundsen Sea during austral summer 2010–2011 and 2014, we collected physical and bio-optical data from ships and autonomous underwater gliders. Gliders documented large phytoplankton blooms associated with Antarctic Surface Waters with low salinity surface water and shallow upper mixed layers (< 50 m). High biomass was not always associated with a specific water mass, suggesting the importance of upper mixed depth and light in influencing phytoplankton biomass. Spectral optical backscatter and ship pigment data suggested that the composition of phytoplankton was spatially heterogeneous, with the large blooms dominated by Phaeocystis and non-bloom waters dominated by diatoms. Phytoplankton growth rates estimated from field data (≤ 0.10 day−1) were at the lower end of the range measured during ship-based incubations, reflecting both in situ nutrient and light limitations. In the bloom waters, phytoplankton biomass was high throughout the 50-m thick upper mixed layer. Those biomass levels, along with the presence of colored dissolved organic matter and detritus, resulted in a euphotic zone that was often < 10 m deep. The net result was that the majority of phytoplankton were light-limited, suggesting that mixing rates within the upper mixed layer were critical to determining the overall productivity; however, regional productivity will ultimately be controlled by water column stability and the depth of the upper mixed layer, which may be enhanced with continued ice melt in the Amundsen Sea Polynya.

We present the first densely-sampled hydrographic survey of the Amundsen Sea Polynya (ASP) region, including a detailed characterization of its freshwater distributions. Multiple components contribute to the freshwater budget, including precipitation, sea ice melt, basal ice shelf melt, and iceberg melt, from local and non-local sources. We used stable oxygen isotope ratios in seawater (δ18O) to distinguish quantitatively the contributions from sea ice and meteoric-derived sources. Meteoric fractions were high throughout the winter mixed layer (WML), with maximum values of 2–3% (±0.5%). Because the ASP region is characterized by deep WMLs, column inventories of total meteoric water were also high, ranging from 10–13 m (±2 m) adjacent to the Dotson Ice Shelf (DIS) and in the deep trough to 7–9 m (±2 m) in shallower areas. These inventories are at least twice those reported for continental shelf waters near the western Antarctic Peninsula. Sea ice melt fractions were mostly negative, indicating net (annual) sea ice formation, consistent with this area being an active polynya. Independently determined fractions of subsurface glacial meltwater (as one component of the total meteoric inventory) had maximum values of 1–2% (±0.5%), with highest and shallowest maximum values at the DIS outflow (80–90 m) and in iceberg-stirred waters (150–200 m). In addition to these upwelling sites, contributions of subsurface glacial meltwater could be traced at depth along the ∼ 27.6 isopycnal, from which it mixes into the WML through various processes. Our results suggest a quasi-continuous supply of melt-laden iron-enriched seawater to the euphotic zone of the ASP and help to explain why the ASP is Antarctica’s most biologically productive polynya per unit area.

Recent attention has focused on accelerated glacial losses along the Amundsen Sea coast that result from changes in atmosphere and ocean circulation, with sea ice playing a mediating but not well-understood role. Here, we investigated how sea ice has changed in the Amundsen Sea over the period of 1979 to 2014, focusing on spatio-temporal changes in ice edge advance/retreat and percent sea ice cover in relation to changes in winds. In contrast to the widespread sea ice decreases to the east and increases to the west of the Amundsen Sea, sea ice changes in the Amundsen Sea were confined to three areas: (i) offshore of the shelf break, (ii) the southern Pine Island Polynya, and (iii) the eastern Amundsen Sea Polynya. Offshore, a 2-month decrease in ice season duration coincided with seasonal shifts in wind speed and direction from March to May (relating to later ice advance) and from September to August (relating to earlier retreat), consistent with reported changes in the depth/location of the Amundsen Sea Low. In contrast, sea ice decreases in the polynya areas corresponded to episodic or step changes in spring ice retreat (earlier by 1–2 months) and were coincident with changes to Thwaites Iceberg Tongue (located between the two polynyas) and increased southeasterly winds. Temporal correlations among these three areas were weak, indicating different local forcing and/or differential response to large-scale forcing. Although our analysis has shown that part of the variability can be explained by changes in winds or to the coastal icescape, an additional but unknown factor is how sea ice has responded to changes in ocean heat and freshwater inputs. Unraveling cause and effect, critical for predicting changes to this rapidly evolving ocean-ice shelf-sea ice system, will require in situ observations, along with improved remote sensing capabilities and ocean modeling.

We report results from a yearlong, moored sediment trap in the Amundsen Sea Polynya (ASP), the first such time series in this remote and productive ecosystem. Results are compared to a long-term (1992–2013) time series from the western Antarctic Peninsula (WAP). The ASP trap was deployed from December 2010 to December 2011 at 350 m depth. We observed two brief, but high flux events, peaking at 8 and 5 mmol C m−2 d−1 in January and December 2011, respectively, with a total annual capture of 315 mmol C m−2. Both peak fluxes and annual capture exceeded the comparable WAP observations. Like the overlying phytoplankton bloom observed during the cruise in the ASP (December 2010 to January 2011), particle flux was dominated by Phaeocystis antarctica, which produced phytodetrital aggregates. Particles at the start of the bloom were highly depleted in 13C, indicating their origin in the cold, CO2-rich winter waters exposed by retreating sea ice. As the bloom progressed, microscope visualization and stable isotopic composition provided evidence for an increasing contribution by zooplankton fecal material. Incubation experiments and zooplankton observations suggested that fecal pellet production likely contributed 10–40% of the total flux during the first flux event, and could be very high during episodic krill swarms. Independent estimates of export from the surface (100 m) were about 5–10 times that captured in the trap at 350 m. Estimated bacterial respiration was sufficient to account for much of the decline in the flux between 50 and 350 m, whereas zooplankton respiration was much lower. The ASP system appears to export only a small fraction of its production deeper than 350 m within the polynya region. The export efficiency was comparable to other polar regions where phytoplankton blooms were not dominated by diatoms.

Special Collection: Special Feature: ASPIRE: The Amundsen Sea Polynya International Research Expedition.-- 17 pages, 4 figures, 7 tables, Despite being perpetually cold, seasonally ice-covered and dark, the coastal Southern Ocean is highly productive and harbors a diverse microbiota. During the austral summer, ice-free coastal patches (or polynyas) form, exposing pelagic organisms to sunlight, triggering intense phytoplankton blooms. This strong seasonality is likely to influence bacterioplankton community composition (BCC). For the most part, we do not fully understand the environmental drivers controlling high-latitude BCC and the biogeochemical cycles they mediate. In this study, the Amundsen Sea Polynya was used as a model system to investigate important environmental factors that shape the coastal Southern Ocean microbiota. Population dynamics in terms of occurrence and activity of abundant taxa was studied in both environmental samples and microcosm experiments by using 454 pyrosequencing of 16S rRNA genes. We found that the BCC in the photic epipelagic zone had low richness, with dominant bacterial populations being related to taxa known to benefit from high organic carbon and nutrient loads (copiotrophs). In contrast, the BCC in deeper mesopelagic water masses had higher richness, featuring taxa known to benefit from low organic carbon and nutrient loads (oligotrophs). Incubation experiments indicated that direct impacts of light and competition for organic nutrients are two important factors shaping BCC in the Amundsen Sea Polynya, The research was funded by the Swedish Research Council (grants to SB) and by the US National Science Foundation Office of Polar Programs (ANT-0839069 to PY)

To evaluate what drives phytoplankton photosynthesis rates in the Amundsen Sea Polynya (ASP), Antarctica, during the spring bloom, we studied phytoplankton biomass, photosynthesis rates, and water column productivity during a bloom of Phaeocystis antarctica (Haptophyceae) and tested effects of iron (Fe) and light availability on these parameters in bioassay experiments in deck incubators. Phytoplankton biomass and productivity were highest (20 µg chlorophyll a L−1 and 6.5 g C m−2 d−1) in the central ASP where sea ice melt water and surface warming enhanced stratification, reducing mixed layer depth and increasing light availability. In contrast, maximum photosynthesis rate (P*max), initial light-limited slope of the photosynthesis–irradiance curve (α*), and maximum photochemical efficiency of photosystem II (Fv/Fm) were highest in the southern ASP near the potential Fe sources of the Dotson and Getz ice shelves. In the central ASP, P*max, α*, and Fv/Fm were all lower. Fe addition increased phytoplankton growth rates in three of twelve incubations, and at a significant level when all experiments were analyzed together, indicating Fe availability may be rate-limiting for phytoplankton growth in several regions of the ASP early in the season during build-up of the spring bloom. Moreover, Fe addition increased P*max, α*, and Fv/Fm in almost all experiments when compared to unamended controls. Incubation under high light also increased P*max, but decreased Fv/Fm and α* when compared to low light incubation. These results indicate that the lower values for P*max, α*, and Fv/Fm in the central ASP, compared to regions close to the ice shelves, are constrained by lower Fe availability rather than light availability. Our study suggests that higher Fe availability (e.g., from higher melt rates of ice shelves) would increase photosynthesis rates in the central ASP and potentially increase water column productivity 1.7-fold, making the ASP even more productive than it is today.

During a springtime study of pelagic microbial activities in the coastal Alaskan Arctic (Chukchi Sea, June 1998), a selective serial dilution technique was used to estimate the relative abundance of cold-loving (psychrophilic) and cold-tolerant (psychrotolerant or psychrotrophic) pelagic bacteria at four stations representing different stages of an algal bloom. Psychrophiles were numerically dominant, regardless of the organic nutrient concentration of their source waters (51–101 µmol L−1 total organic carbon; 2-17 µmol L−1 total organic nitrogen). At all stations, lower temperature incubations generated significantly higher most probable number (MPN) estimates. Further, these psychrophiles were unable to acclimate to 20°C over the course of a 3-month incubation. Our pelagic experiments are the first of their kind reported from the Western Arctic, where surface waters are dominated by the Pacific Ocean inflow, and suggest that psychrophiles can dominate even in waters only recently subjected to perennially cold conditions, independent of nutrient status or bloom condition.

The carbon to nitrogen (C:N) stoichiometry of phytoplankton production varied significantly during the spring–summer bloom in the North Water Polynya (NOW), from April through July 1998. The molar ratio of particulate organic carbon (POC) to nitrogen (PON) production by phytoplankton (ΔPOC:ΔPON) increased from 5.8 during April through early June to 8.9 in late June and July. The molar dissolved inorganic carbon (DIC) to nitrate+nitrite (NO 3 ) drawdown ratio (ΔDIC: ΔNO 3 ) increased from 6.7 in April and May, to 11.9 in June (no estimate for July because of ice melting). The discrepancy between ΔPOC:ΔPON and ΔDIC:ΔNO 3 was likely due to dissolved organic carbon (DOC) production. Increased ΔPOC:ΔPON of phytoplankton and surface water ΔDIC:ΔNO 3 throughout the phytoplankton blooms resulted from changes in physical properties of the upper water column, such as reduced thickness of the surface mixed layer that exposed phytoplankton to increased photosynthetically available radiation (PAR), accompanied by NO 3 depletion. This is expected to have significant effects on the cycling of carbon (C) and nitrogen (N) in pelagic ecosystems, as the increased C:N ratio of organic matter decreases its quality as substrate for grazers and microbial communities. Based on ΔPOC:ΔPON, the ratio of POC to chlorophyll a (Chl) production (ΔPOC:ΔChl) and the relationship between Chl yields and NO 3 depletion, we estimate that 71±17% and 46±20% of the depleted NO 3 went to PON production in the euphotic zone over the polynya from April to early June, and late June to July, respectively. The remaining NO 3 was likely channelled to dissolved organic nitrogen (DON) and heterotrophic bacteria, which were not returned to the dissolved inorganic nitrogen (DIN) pool through recycling during the course of the study. Hence, the autotrophic production of organic N and its recycling by the microbial food web were not coupled temporally.

The Amundsen Sea Polynya (ASP) has, on average, the highest productivity per unit area in Antarctic waters. To investigate community structure and the role that zooplankton may play in utilizing this productivity, animals were collected at six stations inside and outside the ASP using paired “day-night” tows with a 1 m2 MOCNESS. Stations were selected according to productivity based on satellite imagery, distance from the ice edge, and depth of the water column. Depths sampled were stratified from the surface to ∼ 50–100 m above the seafloor. Macrozooplankton were also collected at four stations located in different parts of the ASP using a 2 m2 Metro Net for krill surface trawls (0–120 m). The most abundant groups of zooplankton were copepods, ostracods, and euphausiids. Zooplankton biovolume (0.001 to 1.22 ml m-3) and abundance (0.21 to 97.5 individuals m-3) varied throughout all depth levels, with a midsurface maximum trend at ∼ 60–100 m. A segregation of increasing zooplankton trophic position with depth was observed in the MOCNESS tows. In general, zooplankton abundance was low above the mixed layer depth, a result attributed to a thick layer of the unpalatable colonial haptophyte, Phaeocystis antarctica. Abundances of the ice krill, Euphausia crystallarophias, however, were highest near the edge of the ice sheet within the ASP and larvae:adult ratios correlated with temperature above a depth of 60 m. Total zooplankton abundance correlated positively with chlorophyll a above 150 m, but negative correlations observed for biovolume vs. the proportion of P. antarctica in the phytoplankton estimated from pigment ratios (19’hexanoyloxyfucoxanthin:fucoxanthin) again pointed to avoidance of P. antarctica. Quantifying zooplankton community structure, abundance, and biovolume (biomass) in this highly productive polynya helps shed light on how carbon may be transferred to higher trophic levels and to depth in a region undergoing rapid warming.

The Amundsen Sea Polynya is experiencing large increases in glacial meltwater input and hosts an extremely productive and long-lasting summer phytoplankton bloom, suggesting a crucial role for natural Fe fertilization. Early summer distributions and dynamics of the dissolved bioactive metals Fe, Mn, Zn, Cu and Ni were investigated during a three week period in 2010–2011, using GEOTRACES-compliant methods. Dissolved Fe was very low (0.06–0.12 nmol kg−1) in the upper 20 m of the central polynya, suggesting that the sub-maximal rates of in situ primary productivity reported previously for this growth phase of the bloom are attributable to insufficient Fe availability. Weeks after the sampling period, phytoplankton biomass accumulated to peak bloom conditions, implying a continuous supply of bioavailable Fe to the euphotic zone. The dominant biologically-relevant Fe source was meltwater-enriched seawater flowing from the Dotson Ice Shelf cavity and delivering Fe at 0.7 nmol kg−1 to the broader polynya. The modest Fe content of Circumpolar Deep Water (CDW; 0.3 nmol kg−1), invading through cross-shelf troughs, was strongly augmented by benthic Fe inputs, which may combine with glacial meltwater dFe in the Dotson outflow. Sea ice melting provided a modest local Fe flux, insufficient to drive large annual blooms. Dissolved Mn was strongly reduced in surface waters, but displayed a subsurface maximum likely advected through the region from shallow coastal sediments. Nutrient-type elements Zn, Cu and Ni had large to small dynamic ranges, respectively, and increasing concentrations with depth, indicating uptake and remineralization within the polynya system. Surface water drawdown ratios of metals and nutrients provided novel estimates of metal quotas (metal/P) for the dominant bloom phytoplankton, Phaeocystis antarctica. At one unique mature bloom station, Zn and Cu were scavenged to low concentrations throughout the 350 m water column, a possible result of intense removal onto sinking Phaeocystis biodetritus. The Amundsen Sea appears to be a model region for studying the biogeochemical consequences of increased glacial meltwater inputs.

As part of the first investigation of the North Water region of Baffin Bay to specifically examine carbon cycling in this unique and highly productive area, we found that the distributions of carbon within these waters were controlled by a complex system of transport and biological processes. We systematically collected samples throughout the North Water during April–July 1998 and August–September 1999 and analyzed them for total dissolved inorganic carbon (DIC), alkalinity, dissolved organic carbon, and total suspended particulate carbon. Consistent with biogenic drawdown, surface DIC concentrations dropped by as much as 250 μmol kg−1 during the summer and began to increase again by the end of September. Although the surface waters were supersaturated with carbon dioxide in early spring, extensive ice cover limited CO2 outgassing at that time. As the ice cleared, decreasing surface DIC concentrations supported significant fluxes of CO2 into the ocean. In late September and early October, when ice again was beginning to cover the area, the surface waters were still undersaturated in CO2, implying that the North Water could be a net sink of atmospheric carbon, if winter air–sea fluxes are minimal. There is strong evidence that horizontal advection plays an important role in controlling DIC distributions, although we were unable to independently quantify the advective fluxes. Based on the observed changes in total carbon concentrations and estimates of air–sea fluxes, we found that carbon was lost from the surface waters between April 1998 and October 1999, probably due to both biological and advective export.

Partial pressure of CO2 (pCO2) and dissolved oxygen (DO) in the surface waters of the Amundsen Sea Polynya (ASP) were measured during austral summer 2010–2011 on the Amundsen Sea Polynya International Research Expedition (ASPIRE). Surface pCO2 in the central polynya was as low as 130 µatm, mainly due to strong net primary production. Comparing saturation states of pCO2 and DO distinguished dominant factors (biological activity, temperature, upwelling, and ice melt) controlling pCO2 across regions. Air-sea CO2 flux, estimated using average shipboard winds, showed high spatial variability (-52 to 25 mmol C m-2 d-1) related to these factors. The central region exhibited a high flux of -36 ± 8.4 mmol C m-2 d-1, which is ∼ 50% larger than that reported for the peak of the bloom in the well-studied Ross Sea, comparable to high rates reported for the Chukchi Sea, and significantly higher than reported for most continental shelves around the world. This central region (∼ 20,000 km2) accounted for 85% of the CO2 uptake for the entire open water area. Margins with lower algal biomass accounted for ∼ 15% of regional carbon uptake, likely resulting from pCO2 reductions by sea ice melt. During ASPIRE we also observed pCO2 up to 490 µatm in a small region near the Dotson Ice Shelf with an efflux of 11 ± 5.4 mmol C m-2 d-1 that offset about 3% of the uptake in the much larger central region. Overall, the 2010–2011 ASP was a large net sink for atmospheric CO2 with a spatially averaged flux density of -18 ± 14 mmol C m-2 d-1. This high flux suggests a disproportionate influence on the uptake of CO2 by the Southern Ocean. Since the region has experienced a significant increase in open water duration (1979–2013), we speculate about whether this CO2 sink will increase with future climate-driven change.

Science, Technology, Engineering, and Math (STEM) disciplines have become key focus areas in the education community of the United States. Newly adopted across the nation, Next Generation Science Standards require that educators embrace innovative approaches to teaching. Transforming classrooms to actively engage students through a combination of knowledge and practice develops conceptual understanding and application skills. The partnerships between researchers and educators during the Amundsen Sea Polynya International Research Expedition (ASPIRE) offer an example of how academic research can enhance K-12 student learning. In this commentary, we illustrate how ASPIRE teacher–scientist partnerships helped engage students with actual and virtual authentic scientific investigations. Crosscutting concepts of research in polar marine science can serve as intellectual tools to connect important ideas about ocean and climate science for the public good.

New evidence suggests that cold-loving (psychrophilic) bacteria may be a dynamic component of the episodic bloom events of high-latitude ecosystems. Here we report the results of an unusually early springtime study of pelagic microbial activity in the coastal Alaskan Arctic. Heterotrophic bacterioplankton clearly responded to an algal bloom by doubling cell size, increasing the fraction of actively respiring cells (up to an unprecedented 84% metabolically active using redox dye CTC), shifting substrate-uptake capabilities from kinetic parameters better adapted to lower substrate concentrations to those more suited for higher concentrations, and more than doubling cell abundance. Community composition (determined by polymerase chain reaction/DGGE and nucleotide sequence analysis) also shifted over the bloom. Results support, for the first time with modern molecular methods, previous culture-based observations of bacterial community succession during Arctic algal blooms and confirm that previously observed variability in pelagic microbial activity can be linked to changes in community structure. During early bloom stages, virioplankton and bacterial abundance were comparable, suggesting that mortality due to phage infection was low at that time. The virus-to-bacteria ratio (VBR) increased 10-fold at the height of the bloom, however, suggesting an increased potential for bacterioplankton mortality resulting from viral infection. The peak in VBR coincided with observed shifts in both microbial activity and community structure. These early-season data suggest that substrate and virioplankton interactions may control the active microbial carbon cycling of this region.

To test the hypothesized inhibition of high-latitude marine bacteria living at lower temperatures when organic matter is scarce, the effects of temperature and organic substrate concentration on pelagic microbial heterotrophy were investigated in the perennially cold surface waters of a summertime arctic polynya. Utilization (incorporation plus respiration) of radiolabeled amino acids was measured as a function of increasing added substrate concentration at in situ (subzero) temperature and under short-term warming. Results analyzed using analysis of variance (ANOVA) showed that increased substrate concentration had a significant effect on utilization rates at all stations, suggesting that communities were always living well below saturating levels of substrate. About half of the communities sampled revealed a significant temperature response using ANOVA; interactions between temperature and substrate concentration were detected only rarely. Kinetic parameters, used to link microbial activity and substrate utilization with temperature sensitivity, exhibited mixed responses to short-term warming. Maximum specific utilization rates showed the greater temperature sensitivity, with Q 10 values ranging from 0.25 to 13. Specific affinities responded to temperature significantly at only about half of the stations in the polynya. Psychrophilic behaviors (e.g., highest specific affinities and oligotrophic capacities at lower incubation temperatures) were observed at stations most likely to be influenced by direct Arctic Ocean outflow. Complete agreement with the hypothesis of enhanced substrate requirement by bacteria living at subzero temperature was not found. An improved understanding of the diversity of cold-tolerant and cold-loving microorganisms is needed before generalizing or predicting the role of temperature in the cycling of organic matter at high latitudes.

Chemoautotrophic communities on lipid-rich whale skeletons are known from a total of 16 modern and fossil sites in the deep Pacific Ocean. While the attached fauna of modern whale bones has been studied, the impact of whale falls on surrounding sediment assemblages remains largely unevaluated. Using the research submersible Alvin, we sampled the sediment community at distances of 0, 0.5, 1, 2, 4, and ∼100 m from the lipid-rich skeleton of a 21 m balaenopterid on the 1240 m seafloor in Santa Catalina Basin. When sampled in 1988 and 1991, the skeleton had been on the seafloor for >4 yr and supported a large attached chemoautotrophic assemblage. Sedimentary organic content, microbial biomass and bacterial abundance were not significantly different near the skeleton than in background sediments, and pore-water sulfide concentrations were only modestly elevated (to ⩽20 μM) adjacent to the bones. The species composition of infaunal macrobenthos near the skeleton was similar to that in background sediments, providing little evidence of a specialized enrichment and/or sulfophilic assemblage. Nonetheless, macrofaunal abundance within 0.5 m of the skeleton was reduced by >40%, due to a decline in the paraonid polychaete Levinsenia oculata. The reduction in L. oculata (the community dominant) caused a sharp increase in rarefaction diversity near the skeleton. Bioturbation intensities, evaluated from 234Th profiles, were also dramatically reduced in sediments near the skeleton, as were rates of extracellular lipase activity. We postulate that reduced infaunal abundance and bioturbation near the skeleton resulted from the interference effects of vesicomyid clam-shell debris, and that the low bioturbation rates in turn limited extracellular lipase activity. We conclude that whale skeletons, and the remains of their associated chemoautotrophic assemblages, may physically impact nearby sediment communities for years after the organic and sulfide enrichment effects of whale falls have dissipated, yielding changes in infaunal diversity and bioturbation.

[1] Offshore tropical river plumes are associated with areas of high N2 fixation (diazotrophy) and biological carbon drawdown. Episodic blooms of the diatom Hemiaulus hauckii and its diazotrophic cyanobacterial symbiont Richelia intracellularisare believed to dominate that carbon drawdown, but the mechanism is not well understood. We report primary productivity associated with blooms of these diatom-diazotroph assemblages (DDAs) in the offshore plume of the Amazon River using simultaneous measurements of O2/Ar ratios and the triple-isotope composition of dissolved O2. In these blooms, we observe peaks in net community productivity, but relatively small changes in gross primary productivity, suggesting that DDA blooms increase the ecosystem carbon export ratio more than twofold. These events of enhanced export efficiency lead to biological uptake of dissolved inorganic carbon and silicate, whose longer mixed-layer residence times otherwise obscure the differential impact of DDAs. The shorter-term rate estimates presented here are consistent with the results derived from longer-term geochemical tracers, confirming that DDAs drive a significant biological CO2 pump in tropical oceans.

[1] In austral summer, from December 2008 to January 2009, we investigated the sea-ice carbon dioxide (CO2) system and CO2 controls in the Amundsen and Ross Seas, Antarctica. We sampled seawater, brine and sea ice for the measurements of total alkalinity (AT), total inorganic carbon (DIC), pH, inorganic nutrients, particulate organic carbon (POC) and nitrogen (PON), chlorophyll a, pigments, salinity and temperature. Large variability in all measured parameters was observed in time and space due to the complex sea-ice dynamics. We discuss the controls of the sea-ice CO2 system, such as brine rejection, biological processes, calcium carbonate (CaCO3) precipitation/dissolution and CO2 exchange. Most (80 to 90%) of the DIC loss was due to brine rejection, which suggests that the sea ice acted as an efficient DIC sink from 0.8 and 2.6 mol m−2 yr−1 (9.6–31 g C m−2 yr−1). The remaining change in DIC was to a large extent explained by net biological production. The AT:DIC ratio in the sea ice was higher than in the under-ice water (UIW), with ratios reaching 1.7, which indicated CaCO3 precipitation and concomitant DIC loss in the sea ice. Elevated AT:DIC ratios and carbonate concentrations were also observed in the UIW, which reflect the solid CaCO3 rejected from the ice during melt. The potential for uptake of atmospheric CO2 in the mixed layer increased by approximately 56 μatm due to the combined effect of CaCO3 precipitation during ice formation, and ice melt in summer.

Particle deposition experiments using mimics of biogenous negative relief (“pits”) and low-excess-density particles in a small annular flume indicate a significantly enhanced deposition rate (number of particles per time) compared to smooth, flat patches of the same diameter. This study included flow visualizations as well as observations of particle residence times, particle concentrations in the pits, and particle fluxes to the pits from the main flow. Experimental conditions of particle concentration, shear velocity, and particle settling velocity mimicked the dynamic characteristics (low excess density and large size) of organic-rich floes and flow conditions in the subtidal and deep sea where biogenous pits are common features. Results suggest that pits provide benthic organisms an important capture mechanism for such floes. Flow visualizations concur qualitatively with previously reported results for twodimensional cavity flow, with unique features due to the conical shape of the pits. When the Rouse number (settling velocity/shear velocity) was much less than 1, pit deposition rate increased with increasing pit aspect ratio (AR = depth/diameter; ranging from 0.25 to 2) and always exceeded deposition to a flat patch of comparable diameter. For the single aspect ratio tested (AR = 0.5) under conditions of increasing turbulence, deposition to the pit increased under transitional flow, but then decreased to near zero when conditions reached fully rough flow. Relative enhancement of deposition to this pit decreased with increased ambient bed roughness since grave1 beds also effectively collect particles. Particle concentration inside pits decreased weakly with pit aspect ratio but greatly increased with increasing roughness Reynolds number. Particle residence time increased somewhat with pit aspect ratio but decreased significantly with increasing roughness Reynolds number. Particle flux into pits from the main flow increased with both increasing aspect ratio and increasing roughness Reynolds number. Enhancement of food supply to pit inhabitants thus depends on the flow regime.