Nutrients and Other Environmental Factors Influence Virus Abundances across Oxic and Hypoxic Marine Environments.

Virus particles are highly abundant in seawater and, on average, outnumber microbial cells approximately 10-fold at the surface and 16-fold in deeper waters; yet, this relationship varies across environments. Here, we examine the influence of a suite of environmental variables, including nutrient concentrations, salinity and temperature, on the relationship between the abundances of viruses and prokaryotes over a broad range of spatial and temporal scales, including along a track from the Northwest Atlantic to the Northeast Pacific via the Arctic Ocean, and in the coastal waters of British Columbia, Canada. Models of varying complexity were tested and compared for best fit with the Akaike Information Criterion, and revealed that nitrogen and phosphorus concentrations, as well as prokaryote abundances, either individually or combined, had significant effects on viral abundances in all but hypoxic environments, which were only explained by a combination of physical and chemical factors. Nonetheless, multivariate models of environmental variables showed high explanatory power, matching or surpassing that of prokaryote abundance alone. Incorporating both environmental variables and prokaryote abundances into multivariate models significantly improved the explanatory power of the models, except in hypoxic environments. These findings demonstrate that environmental factors could be as important as, or even more important than, prokaryote abundance in describing viral abundance across wide-ranging marine environments.

Greater role for Atlantic inflows on sea-ice loss in the Eurasian Basin of the Arctic Ocean.

Arctic sea-ice loss is a leading indicator of climate change and can be attributed, in large part, to atmospheric forcing. Here, we show that recent ice reductions, weakening of the halocline, and shoaling of the intermediate-depth Atlantic Water layer in the eastern Eurasian Basin have increased winter ventilation in the ocean interior, making this region structurally similar to that of the western Eurasian Basin. The associated enhanced release of oceanic heat has reduced winter sea-ice formation at a rate now comparable to losses from atmospheric thermodynamic forcing, thus explaining the recent reduction in sea-ice cover in the eastern Eurasian Basin. This encroaching "atlantification" of the Eurasian Basin represents an essential step toward a new Arctic climate state, with a substantially greater role for Atlantic inflows.

Effect of introducing a competitor on cyclic dominance of sockeye salmon.

We study the effects of introducing a competing species into a 3-species model for the population dynamics of sockeye salmon, thereby converting a food chain into a diamond module. We find that this often leads to the disappearance of the 4-year oscillation of sockeye salmon known as cyclic dominance when parameters are chosen such that all four species can coexist. Only when the population size of the competitor is small the phenomenon of cyclic dominance can persist. There is also a large region of parameter space where either the sockeye salmon or the competitor goes extinct. We discuss how these findings can be reconciled with the prevalence of cyclic dominance in many sockeye brood lakes.

Detecting and coping with disruptive shocks in Arctic marine systems: a resilience approach to place and people.

It seems inevitable that the ongoing and rapid changes in the physical environment of the marine Arctic will push components of the region's existing social-ecological systems-small and large-beyond tipping points and into new regimes. Ongoing changes include warming, freshening, acidification, and alterations to food web structure. In anticipation we pose three distinct but interrelated challenges: (1) to explore existing connectivities within components of the marine system; (2) to seek indicators (if they exist) of approaching regime change through observation and modeling; and (3) to build functional resilience into existing systems through adaptation-oriented policy and to have in hand transformative options when tipping points are crossed and new development trajectories are required. Each of the above challenges is scale dependent, and each requires a much deeper understanding than we currently have of connectivity within existing systems and their response to external forcing. Here, we argue from a global perspective the need to understand the Arctic's role in an increasingly nonlinear world; then describe emerging evidence from new observations on the connectivity of processes and system components from the Canada Basin and subarctic seas surrounding northern North America; and finally posit an approach founded in "resilience thinking" to allow northern residents living in small coastal communities to participate in the observation, adaption and-if necessary-transformation of the social-ecological system with which they live.

Arctic Ocean microbial community structure before and after the 2007 record sea ice minimum.

Increasing global temperatures are having a profound impact in the Arctic, including the dramatic loss of multiyear sea ice in 2007 that has continued to the present. The majority of life in the Arctic is microbial and the consequences of climate-mediated changes on microbial marine food webs, which are responsible for biogeochemical cycling and support higher trophic levels, are unknown. We examined microbial communities over time by using high-throughput sequencing of microbial DNA collected between 2003 and 2010 from the subsurface chlorophyll maximum (SCM) layer of the Beaufort Sea (Canadian Arctic). We found that overall this layer has freshened and concentrations of nitrate, the limiting nutrient for photosynthetic production in Arctic seas, have decreased. We compared microbial communities from before and after the record September 2007 sea ice minimum and detected significant differences in communities from all three domains of life. In particular, there were significant changes in species composition of Eukarya, with ciliates becoming more common and heterotrophic marine stramenopiles (MASTs) accounting for a smaller proportion of sequences retrieved after 2007. Within the Archaea, Marine Group I Thaumarchaeota, which earlier represented up to 60% of the Archaea sequences in this layer, have declined to <10%. Bacterial communities overall were less diverse after 2007, with a significant decrease of the Bacteroidetes. These significant shifts suggest that the microbial food webs are sensitive to physical oceanographic changes such as those occurring in the Canadian Arctic over the past decade.

A three-species model explaining cyclic dominance of Pacific salmon.

The four-year oscillations of the number of spawning sockeye salmon (Oncorhynchus nerka) that return to their native stream within the Fraser River basin in Canada are a striking example of population oscillations. The period of the oscillation corresponds to the dominant generation time of these fish. Various-not fully convincing-explanations for these oscillations have been proposed, including stochastic influences, depensatory fishing, or genetic effects. Here, we show that the oscillations can be explained as an attractor of the population dynamics, resulting from a strong resonance near a Neimark Sacker bifurcation. This explains not only the long-term persistence of these oscillations, but also reproduces correctly the empirical sequence of salmon abundance within one period of the oscillations. Furthermore, it explains the observation that these oscillations occur only in sockeye stocks originating from large oligotrophic lakes, and that they are usually not observed in salmon species that have a longer generation time.

Aragonite undersaturation in the Arctic Ocean: effects of ocean acidification and sea ice melt.

The increase in anthropogenic carbon dioxide emissions and attendant increase in ocean acidification and sea ice melt act together to decrease the saturation state of calcium carbonate in the Canada Basin of the Arctic Ocean. In 2008, surface waters were undersaturated with respect to aragonite, a relatively soluble form of calcium carbonate found in plankton and invertebrates. Undersaturation was found to be a direct consequence of the recent extensive melting of sea ice in the Canada Basin. In addition, the retreat of the ice edge well past the shelf-break has produced conditions favorable to enhanced upwelling of subsurface, aragonite-undersaturated water onto the Arctic continental shelf. Undersaturation will affect both planktonic and benthic calcifying biota and therefore the composition of the Arctic ecosystem.

Smallest algae thrive as the Arctic Ocean freshens.

As climate changes and the upper Arctic Ocean receives more heat and fresh water, it becomes more difficult for mixing processes to deliver nutrients from depth to the surface for phytoplankton growth. Competitive advantage will presumably accrue to small cells because they are more effective in acquiring nutrients and less susceptible to gravitational settling than large cells. Since 2004, we have discerned an increase in the smallest algae and bacteria along with a concomitant decrease in somewhat larger algae. If this trend toward a community of smaller cells is sustained, it may lead to reduced biological production at higher trophic levels.

Archaeal diversity and a gene for ammonia oxidation are coupled to oceanic circulation.

Evidence of microbial zonation in the open ocean is rapidly accumulating, but while the distribution of communities is often described according to depth, the other physical factors structuring microbial diversity and function remain poorly understood. Here we identify three different water masses in the North Water (eastern Canadian Arctic), defined by distinct temperature and salinity characteristics, and show that they contained distinct archaeal communities. Moreover, we found that one of the water masses contained an increased abundance of the archaeal alpha-subunit of the ammonia monooxygenase gene (amoA) and accounted for 70% of the amoA gene detected overall. This indicates likely differences in putative biogeochemical capacities among different water masses. The ensemble of our results strongly suggest that the widely accepted view of depth stratification did not explain microbial diversity, but rather that parent water masses provide the framework for predicting communities and potential microbial function in an Arctic marine system. Our results emphasize that microbial distributions are strongly influenced by oceanic circulation, implying that shifting currents and water mass boundaries resulting from climate change may well impact patterns of microbial diversity by displacing whole biomes from their historic distributions. This relocation could have the potential to establish a substantially different geography of microbial-driven biogeochemical processes and associated oceanic production.

A major ecosystem shift in the northern Bering Sea.

Until recently, northern Bering Sea ecosystems were characterized by extensive seasonal sea ice cover, high water column and sediment carbon production, and tight pelagic-benthic coupling of organic production. Here, we show that these ecosystems are shifting away from these characteristics. Changes in biological communities are contemporaneous with shifts in regional atmospheric and hydrographic forcing. In the past decade, geographic displacement of marine mammal population distributions has coincided with a reduction of benthic prey populations, an increase in pelagic fish, a reduction in sea ice, and an increase in air and ocean temperatures. These changes now observed on the shallow shelf of the northern Bering Sea should be expected to affect a much broader portion of the Pacific-influenced sector of the Arctic Ocean.