Warming and CO2 Enhance Arctic Heterotrophic Microbial Activity

Ocean acidification and warming are two main consequences of climate change that can directly affect biological and ecosystem processes in marine habitats. The Arctic Ocean is the region of the world experiencing climate change at the steepest rate compared with other latitudes. Since marine planktonic microorganisms play a key role in the biogeochemical cycles in the ocean it is crucial to simultaneously evaluate the effect of warming and increasing CO2 on marine microbial communities. In 20 L experimental microcosms filled with water from a high-Arctic fjord (Svalbard), we examined changes in phototrophic and heterotrophic microbial abundances and processes [bacterial production (BP) and mortality], and viral activity (lytic and lysogenic) in relation to warming and elevated CO2. The summer microbial plankton community living at 1.4°C in situ temperature, was exposed to increased CO2 concentrations (135–2,318 μatm) in three controlled temperature treatments (1, 6, and 10°C) at the UNIS installations in Longyearbyen (Svalbard), in summer 2010. Results showed that chlorophyll a concentration decreased at increasing temperatures, while BP significantly increased with pCO2 at 6 and 10°C. Lytic viral production was not affected by changes in pCO2 and temperature, while lysogeny increased significantly at increasing levels of pCO2, especially at 10°C (R2 = 0.858, p = 0.02). Moreover, protistan grazing rates showed a positive interaction between pCO2 and temperature. The averaged percentage of bacteria grazed per day was higher (19.56 ± 2.77% d-1) than the averaged percentage of lysed bacteria by virus (7.18 ± 1.50% d-1) for all treatments. Furthermore, the relationship among microbial abundances and processes showed that BP was significantly related to phototrophic pico/nanoflagellate abundance in the 1°C and the 6°C treatments, and BP triggered viral activity, mainly lysogeny at 6 and 10°C, while bacterial mortality rates was significantly related to bacterial abundances at 6°C. Consequently, our experimental results suggested that future increases in water temperature and pCO2 in Arctic waters will produce a decrease of phytoplankton biomass, enhancement of BP and changes in the carbon fluxes within the microbial food web. All these heterotrophic processes will contribute to weakening the CO2 sink capacity of the Arctic plankton community.En prens

Effects of elevated CO2 on phytoplankton community biomass and species composition during a spring Phaeocystis spp. bloom in the western English Channel

A 21-year time series of phytoplankton community structure was analysed in relation to Phaeocystis spp. to elucidate its contribution to the annual carbon budget at station L4 in the western English Channel (WEC). Between 1993–2014 Phaeocystis spp. contributed ∼4.6% of the annual phytoplankton carbon and during the March − May spring bloom, the mean Phaeocystis spp. biomass constituted 17% with a maximal contribution of 47% in 2001. Upper maximal weekly values above the time series mean ranged from 63 to 82% of the total phytoplankton carbon (∼42–137 mg carbon (C) m −3 ) with significant inter-annual variability in Phaeocystis spp. Maximal biomass usually occurred by the end of April, although in some cases as early as mid-April (2007) and as late as late May (2013). The effects of elevated pCO 2 on the Phaeocystis spp. spring bloom were investigated during a fifteen-day semi-continuous microcosm experiment. The phytoplankton community biomass was estimated at ∼160 mg C m −3 and was dominated by nanophytoplankton (40%, excluding Phaeocystis spp.), Phaeocystis spp. (30%) and cryptophytes (12%). The smaller fraction of the community biomass comprised picophytoplankton (9%), coccolithophores (3%), Synechococcus (3%), dinoflagellates (1.5%), ciliates (1%) and diatoms (0.5%). Over the experimental period, total biomass increased significantly by 90% to ∼305 mg C m −3 in the high CO 2 treatment while the ambient pCO 2 control showed no net gains. Phaeocystis spp. exhibited the greatest response to the high CO 2 treatment, increasing by 330%, from ∼50 mg C m −3 to over 200 mg C m −3 and contributing ∼70% of the total biomass. Taken together, the results of our microcosm experiment and analysis of the time series suggest that a future high CO 2 scenario may favour dominance of Phaeocystis spp. during the spring bloom. This has significant implications for the formation of hypoxic zones and the alteration of food web structure including inhibitory feeding effects and lowered fecundity in many copepod species

Net primary productivity estimates and environmental variables in the Arctic Ocean: An assessment of coupled physical-biogeochemical models

The relative skill of 21 regional and global biogeochemical models was assessed in terms of how well the models reproduced observed net primary productivity (NPP) and environmental variables such as nitrate concentration (NO3), mixed layer depth (MLD), euphotic layer depth (Zeu), and sea ice concentration, by comparing results against a newly updated, quality-controlled in situ NPP database for the Arctic Ocean (1959-2011). The models broadly captured the spatial features of integrated NPP (iNPP) on a pan-Arctic scale. Most models underestimated iNPP by varying degrees in spite of overestimating surface NO3, MLD, and Zeu throughout the regions. Among the models, iNPP exhibited little difference over sea ice condition (ice-free vs. ice-influenced) and bottom depth (shelf vs. deep ocean). The models performed relatively well for the most recent decade and towards the end of Arctic summer. In the Barents and Greenland Seas, regional model skill of surface NO3 was best associated with how well MLD was reproduced. . Regionally, iNPP was relatively well simulated in the Beaufort Sea and the central Arctic Basin, where in situ NPP is low and nutrients are mostly depleted. Models performed less well at simulating iNPP in the Greenland and Chukchi Seas, despite the higher model skill in MLD and sea ice concentration, respectively. iNPP model skill was constrained by different factors in different Arctic Ocean regions. Our study suggests that better parameterization of biological and ecological microbial rates (phytoplankton growth and zooplankton grazing) are needed for improved Arctic Ocean biogeochemical modeling

Thermal thresholds of phytoplankton growth in polar waters and their consequences for a warming polar ocean

Polar areas are experiencing the steepest warming rates on Earth, a trend expected to continue in the future. In these habitats, phytoplankton communities constitute the basis of the food web and their thermal tolerance may dictate how warming affects these delicate environments. Here, we compiled available data on thermal responses of phytoplankton growth in polar waters. We assembled 53 growth-vs.-temperature curves (25 from the Arctic, 28 from the Southern oceans), indicating the limited information available for these ecosystems. Half of the data from Arctic phytoplankton came from natural communities where low ambient concentrations could limit growth rates. Phytoplankton from polar waters grew faster under small temperature increases until reaching an optimum (TOPT), and slowed when temperatures increased beyond this value. This left-skewed curves were characterized by higher activation energies (Ea) for phytoplankton growth above than below the TOPT. Combining these thermal responses we obtained a community TOPT of 6.5◦C (±0.2) and 5.2◦C (±0.1) for Arctic and Southern Ocean phytoplankton communities, respectively. These threshold temperatures were already exceeded at 70◦N during the first half of August 2013, evidenced by sea surface temperatures (SSTs, satellite data, http://www.ncdc.noaa.gov). We forecasted SSTs for the end of the twenty-first century by assuming an overall 3◦C increase, equivalent to a low emission scenario. Our forecasts show that SSTs at 70◦N are expected to exceed TOPT during summer by 2100, and during the first half of August at 75◦N. While recent Arctic spring temperatures average 0.5◦C and −0.7◦C at 70◦N and 75◦N, respectively, they could increase to 2.8◦C at 70◦N and 2.2◦C at 75◦N as we approach 2100. Such temperature increases could lead to intense phytoplankton blooms, shortened by fast nutrient consumption. As SSTs increase, thermal thresholds for phytoplankton growth would be eventually exceeded during bloom development. This could lead to changes in the blooming phytoplankton community, threatening the production peak and cycles in the Arctic. Our forecasted phytoplankton responses, are constrained by the limited data set, besides uncertainties in the most plausible future Arctic temperature scenarios. To improve predictions in polar oceans, we need to increase the number of studies, in particular for a fast-changing Arctic

Thermal Thresholds of Phytoplankton Growth in Polar Waters and Their Consequences for a Warming Polar Ocean

Polar areas are experiencing the steepest warming rates on Earth, a trend expected to continue in the future. In these habitats, phytoplankton communities constitute the basis of the food web and their thermal tolerance may dictate how warming affects these delicate environments. Here, we compiled available data on thermal responses of phytoplankton growth in polar waters. We assembled 53 growth-vs.-temperature curves (25 from the Arctic, 28 from the Southern oceans), indicating the limited information available for these ecosystems. Half of the data from Arctic phytoplankton came from natural communities where low ambient concentrations could limit growth rates. Phytoplankton from polar waters grew faster under small temperature increases until reaching an optimum (TOPT), and slowed when temperatures increased beyond this value. This left-skewed curves were characterized by higher activation energies (Ea) for phytoplankton growth above than below the TOPT. Combining these thermal responses we obtained a community TOPT of 6.5°C (±0.2) and 5.2°C (±0.1) for Arctic and Southern Ocean phytoplankton communities, respectively. These threshold temperatures were already exceeded at 70°N during the first half of August 2013, evidenced by sea surface temperatures (SSTs, satellite data, http://www.ncdc.noaa.gov). We forecasted SSTs for the end of the twenty-first century by assuming an overall 3°C increase, equivalent to a low emission scenario. Our forecasts show that SSTs at 70°N are expected to exceed TOPT during summer by 2100, and during the first half of August at 75°N. While recent Arctic spring temperatures average 0.5°C and −0.7°C at 70°N and 75°N, respectively, they could increase to 2.8°C at 70°N and 2.2°C at 75°N as we approach 2100. Such temperature increases could lead to intense phytoplankton blooms, shortened by fast nutrient consumption. As SSTs increase, thermal thresholds for phytoplankton growth would be eventually exceeded during bloom development. This could lead to changes in the blooming phytoplankton community, threatening the production peak and cycles in the Arctic. Our forecasted phytoplankton responses, are constrained by the limited data set, besides uncertainties in the most plausible future Arctic temperature scenarios. To improve predictions in polar oceans, we need to increase the number of studies, in particular for a fast-changing Arctic

Design and use of a new primer pair for the characterization of the cyanobacteria Synechococcus and Prochlorococcus communities targeting petB gene through metabarcoding approaches

During the last years, the application of next-generation sequencing (NGS) technologies to search for specific genetic markers has become a crucial method for the characterization of microbial communities. Illumina MiSeq, likely the most widespread NGS platform for metabarcoding experiments and taxonomic classification, allows processing shorter reads than the classical SANGER sequencing method and therefore requires specific primer pairs that produce shorter amplicons. Specifically, for the analysis of the commonly studied Prochlorococcus and Synechococcus communities, the petB marker gene has recently stood out as able to provide deep coverage to determine the microdiversity of the community. However, current petB primer set produce a 597 bp amplicon that is not suitable for MiSeq chemistry.Here, we designed and tested a petB primer pair that targets both Prochlorococcus and Synechococcus communities producing an appropriate amplicon to be used with state-of-the-art Illumina MiSeq. This new primer set allows the classification of both groups to a low taxonomic level and is therefore suitable for high throughput experiments using MiSeq technologies, therefore constituting a useful, novel tool to facilitate further studies on Prochlorococcus and Synechococcus communities. • This work describes the de novo design of a Prochlorococcus and Synechococcus-specific petB primer pair, allowing the characterization of both populations to a low taxonomic level. • This primer pair is suitable for widespread Illumina MiSeq sequencing technologies. • petB was confirmed as an adequate target for the characterization of both picocyanobacteria

Zooplankton excretion metabolites stimulate Southern Ocean phytoplankton growth

Warming over Antarctica is leading to changes in the zooplankton communities inhabiting the Southern Ocean. It has been observed that zooplankton not only regulates phytoplankton through grazing, but also through the recycling of nutrients that are essential for phytoplankton growth. In this way, the effects of warming on zooplankton populations will change the amount or proportion at which recycled nutrients are restored. To estimate how the recycled nutrients released by zooplankton populations, dominated by krill (Euphausia superba), amphipods or copepods, affect the phytoplankton uptake and communities, we performed four incubation experiments: two close to the Antarctic Peninsula and two at the Southern Atlantic Ocean. Our results showed a stimulating effect of the addition of metabolites on ammonia removal rates and on the net growth of phytoplankton communities, with different responses amongst the different phytoplankton groups. According to our results, phytoplankton net growth and community composition may be altered if this relevant source of nutrients is lost due to projected changes in the abundance or distribution of these zooplankton populations

Interactive effect of temperature and CO2 increase in Arctic phytoplankton

An experiment was performed in order to analyze the effects of the increase in water temperature and CO2 partial pressure expected for the end of this century in a present phytoplankton community inhabiting the Arctic Ocean. We analyzed both factors acting independently and together, to test possible interactions between them. The arctic planktonic community was incubated under six different treatments combining three experimental temperatures (1, 6, and 10°C) with two different CO2 levels of 380 or 1000 ppm, at the UNIS installations in Longyearbyen (Svalbard), in summer 2010. Under warmer temperatures, a decrease in chlorophyll a concentration, biovolume and primary production was found, together with a shift in community structure toward a dominance of smaller cells (nano-sized). Effects of increased pCO2 were more modest, and although interactions were weak, our results suggest antagonistic interactive effects amongst increased temperature and CO2 levels, as elevated CO2 compensated partially the decrease in phytoplankton biomass induced by temperature in some groups. Interactions between the two stressors were generally weak, but elevated CO2 was observed to lead to a steeper decline in primary production with warming. Our results also suggest that future increases in water temperature and pCO2 would lead to a decrease in the community chl a concentration and biomass in the Arctic phytoplankton communities examined, leading to communities dominated by smaller nano-phytoplankton groups, with important consequences for the flow of carbon and food web dynamics.This study was supported by the project Arctic Tipping Points (ATP, contract # 226248) from the European Union. Alexandra Coello-Camba was supported by a grant BES-2007-15193 from the Spanish Ministry of Science and Innovation.Peer reviewedPeer Reviewe

Experimentally determined temperature thresholds for Arctic plankton community metabolism

Climate warming is especially severe in the Arctic, where the average temperature is increasing 0.4 C per decade, two to three times higher than the global average rate. Furthermore, the Arctic has lost more than half of its summer ice extent since 1980 and predictions suggest that the Arctic will be ice free in the summer as early as 2050, which could increase the rate of warming. Predictions based on the metabolic theory of ecology assume that temperature increase will enhance metabolic rates and thus both the rate of primary production and respiration will increase. However, these predictions do not consider the specific metabolic balance of the communities. We tested, experimentally, the response of Arctic plankton communities to seawater temperature spanning from 1 C to 10 C. Two types of communities were tested, open-ocean Arctic communities from water collected in the Barents Sea and Atlantic influenced fjord communities from water collected in the Svalbard fjord system. Metabolic rates did indeed increase as suggested by metabolic theory, however these results suggest an experimental temperature threshold of 5 C, beyond which the metabolism of plankton communities shifts from autotrophic to heterotrophic. This threshold is also validated by field measurements across a range of temperatures which suggested a temperature 5.4 C beyond which Arctic plankton communities switch to heterotrophy. Barents Sea communities showed a much clearer threshold response to temperature manipulations than fjord communities. © Author(s) 2013.Peer Reviewe

Experimentally determined temperature thresholds for Arctic plankton community metabolism.

Climate warming is especially severe in the Arctic, where the average temperature is increasing 0.4 degrees C per decade, two to three times higher than the global average rate. Furthermore, the Arctic has lost more than half of its summer ice extent since 1980 and predictions suggest that the Arctic will be ice free in the summer as early as 2050, which could increase the rate of warming. Predictions based on the metabolic theory of ecology assume that temperature increase will enhance metabolic rates and thus both the rate of primary production and respiration will increase. However, these predictions do not consider the specific metabolic balance of the communities. We tested, experimentally, the response of Arctic plankton communities to seawater temperature spanning from 1 degrees C to 10 degrees C. Two types of communities were tested, open-ocean Arctic communities from water collected in the Barents Sea and Atlantic influenced fjord communities from water collected in the Svalbard fjord system. Metabolic rates did indeed increase as suggested by metabolic theory, however these results suggest an experimental temperature threshold of 5 degrees C, beyond which the metabolism of plankton communities shifts from autotrophic to heterotrophic. This threshold is also validated by field measurements across a range of temperatures which suggested a temperature 5.4 degrees C beyond which Arctic plankton communities switch to heterotrophy. Barents Sea communities showed a much clearer threshold response to temperature manipulations than fjord communities