The Outsized Role of Salps in Carbon Export in the Subarctic Northeast Pacific Ocean.

Periodic blooms of salps (pelagic tunicates) can result in high export of organic matter, leading to an "outsized" role in the ocean's biological carbon pump (BCP). However, due to their episodic and patchy nature, salp blooms often go undetected and are rarely included in measurements or models of the BCP. We quantified salp-mediated export processes in the northeast subarctic Pacific Ocean in summer of 2018 during a bloom of Salpa aspera. Salps migrated from 300 to 750 m during the day into the upper 100 m at night. Salp fecal pellet production comprised up to 82% of the particulate organic carbon (POC) produced as fecal pellets by the entire epipelagic zooplankton community. Rapid sinking velocities of salp pellets (400-1,200 m d-1) and low microbial respiration rates on pellets (<1% of pellet C respired day-1) led to high salp pellet POC export from the euphotic zone-up to 48% of total sinking POC across the 100 m depth horizon. Salp active transport of carbon by diel vertical migration and carbon export from sinking salp carcasses was usually <10% of the total sinking POC flux. Salp-mediated export markedly increased BCP efficiency, increasing by 1.5-fold the proportion of net primary production exported as POC across the base of the euphotic zone and by 2.6-fold the proportion of this POC flux persisting 100 m below the euphotic zone. Salps have unique and important effects on ocean biogeochemistry and, especially in low flux settings, can dramatically increase BCP efficiency and thus carbon sequestration.

Interactive Effects of CO2 , Temperature, Irradiance, and Nutrient Limitation on the Growth and Physiology of the Marine Diatom Thalassiosira pseudonana (Coscinodiscophyceae).

The marine diatom Thalassiosira pseudonana was grown in continuous culture systems to study the interactive effects of temperature, irradiance, nutrient limitation, and the partial pressure of CO2 (pCO2 ) on its growth and physiological characteristics. The cells were able to grow at all combinations of low and high irradiance (50 and 300 µmol photons · m-2  · s-1 , respectively, of visible light), low and high pCO2 (400 and 1,000 µatm, respectively), nutrient limitation (nitrate-limited and nutrient-replete conditions), and temperatures of 10-32°C. Under nutrient-replete conditions, there was no adverse effect of high pCO2 on growth rates at temperatures of 10-25°C. The response of the cells to high pCO2 was similar at low and high irradiance. At supraoptimal temperatures of 30°C or higher, high pCO2 depressed growth rates at both low and high irradiance. Under nitrate-limited conditions, cells were grown at 38 ± 2.4% of their nutrient-saturated rates at the same temperature, irradiance, and pCO2 . Dark respiration rates consistently removed a higher percentage of production under nitrate-limited versus nutrient-replete conditions. The percentages of production lost to dark respiration were positively correlated with temperature under nitrate-limited conditions, but there was no analogous correlation under nutrient-replete conditions. The results suggest that warmer temperatures and associated more intense thermal stratification of ocean surface waters could lower net photosynthetic rates if the stratification leads to a reduction in the relative growth rates of marine phytoplankton, and at truly supraoptimal temperatures there would likely be a synergistic interaction between the stresses from temperature and high pCO2 (lower pH).

Sustained deposition of contaminants from the Deepwater Horizon spill.

The 2010 Deepwater Horizon oil spill resulted in 1.6-2.6 × 10(10) grams of petrocarbon accumulation on the seafloor. Data from a deep sediment trap, deployed 7.4 km SW of the well between August 2010 and October 2011, disclose that the sinking of spill-associated substances, mediated by marine particles, especially phytoplankton, continued at least 5 mo following the capping of the well. In August/September 2010, an exceptionally large diatom bloom sedimentation event coincided with elevated sinking rates of oil-derived hydrocarbons, black carbon, and two key components of drilling mud, barium and olefins. Barium remained in the water column for months and even entered pelagic food webs. Both saturated and polycyclic aromatic hydrocarbon source indicators corroborate a predominant contribution of crude oil to the sinking hydrocarbons. Cosedimentation with diatoms accumulated contaminants that were dispersed in the water column and transported them downward, where they were concentrated into the upper centimeters of the seafloor, potentially leading to sustained impact on benthic ecosystems.

Experimental strategies to assess the biological ramifications of multiple drivers of global ocean change-A review.

Marine life is controlled by multiple physical and chemical drivers and by diverse ecological processes. Many of these oceanic properties are being altered by climate change and other anthropogenic pressures. Hence, identifying the influences of multifaceted ocean change, from local to global scales, is a complex task. To guide policy-making and make projections of the future of the marine biosphere, it is essential to understand biological responses at physiological, evolutionary and ecological levels. Here, we contrast and compare different approaches to multiple driver experiments that aim to elucidate biological responses to a complex matrix of ocean global change. We present the benefits and the challenges of each approach with a focus on marine research, and guidelines to navigate through these different categories to help identify strategies that might best address research questions in fundamental physiology, experimental evolutionary biology and community ecology. Our review reveals that the field of multiple driver research is being pulled in complementary directions: the need for reductionist approaches to obtain process-oriented, mechanistic understanding and a requirement to quantify responses to projected future scenarios of ocean change. We conclude the review with recommendations on how best to align different experimental approaches to contribute fundamental information needed for science-based policy formulation.

Chemical dispersants can suppress the activity of natural oil-degrading microorganisms.

During the Deepwater Horizon oil well blowout in the Gulf of Mexico, the application of 7 million liters of chemical dispersants aimed to stimulate microbial crude oil degradation by increasing the bioavailability of oil compounds. However, the effects of dispersants on oil biodegradation rates are debated. In laboratory experiments, we simulated environmental conditions comparable to the hydrocarbon-rich, 1,100 m deep plume that formed during the Deepwater Horizon discharge. The presence of dispersant significantly altered the microbial community composition through selection for potential dispersant-degrading Colwellia, which also bloomed in situ in Gulf deep waters during the discharge. In contrast, oil addition to deepwater samples in the absence of dispersant stimulated growth of natural hydrocarbon-degrading Marinobacter. In these deepwater microcosm experiments, dispersants did not enhance heterotrophic microbial activity or hydrocarbon oxidation rates. An experiment with surface seawater from an anthropogenically derived oil slick corroborated the deepwater microcosm results as inhibition of hydrocarbon turnover was observed in the presence of dispersants, suggesting that the microcosm findings are broadly applicable across marine habitats. Extrapolating this comprehensive dataset to real world scenarios questions whether dispersants stimulate microbial oil degradation in deep ocean waters and instead highlights that dispersants can exert a negative effect on microbial hydrocarbon degradation rates.

Transparent exopolymer particles: from aquatic environments and engineered systems to membrane biofouling.

Transparent exopolymer particles (TEP) are ubiquitous in marine and freshwater environments. For the past two decades, the distribution and ecological roles of these polysaccharide microgels in aquatic systems were extensively investigated. More recent studies have implicated TEP as an active agent in biofilm formation and membrane fouling. Since biofouling is one of the main hurdles for efficient operation of membrane-based technologies, there is a heightened interest in understanding the role of TEP in engineered water systems. In this review, we describe relevant TEP terminologies while critically discussing TEP biological origin, biochemical and physical characteristics, and occurrence and distributions in aquatic systems. Moreover, we examine the contribution of TEP to biofouling of various membrane technologies used in the desalination and water/wastewater treatment industry. Emphasis is given to the link between TEP physicochemical and biological properties and the underlying biofouling mechanisms. We highlight that thorough understanding of TEP dynamics in feedwater sources, pretreatment challenges, and biofouling mechanisms will lead to better management of fouling/biofouling in membrane technologies.

Why biofouling cannot contribute to the vertical transport of small microplastic.

In contrast to expectations, even buoyant microplastics like polyethylene and polypropylene are found at high concentrations in deep sediment traps and deep-sea sediments. To explain the presence of such buoyant microplastic particles at great ocean depths, several vertical transport mechanisms are under discussion with biofouling as one of the most referred. Biofouling is thought to increase the density of microplastic particles to the point that they sink to the deep sea, but this has mostly been shown on large microplastic particles ≥ 1 mm. However, although microplastics are defined as particles between 1 and 5000 µm, most microplastics are < 100 µm. In the ocean plastic particles continuously fragment, converting each "large" particle into several "small" particles, and particle abundance increases drastically with decreasing size. We argue that biofouling is not a reasonable transport mechanism for small microplastic particles ≤ 100 µm, which form the majority of microplastics. Biofilm density depends on its community and composition. A biofilm matrix of extracellular polymeric substances and bacteria has a lower density than seawater, in contrast to diatoms or large organisms like mussels or barnacles. We suggest that a small microplastic particle cannot host a biofilm community consisting of the heavy organisms required to induce sinking. Furthermore, to reach the deep sea within a reasonable timespan, a microplastic particle needs to sink several meters per day. Therefore, the excess density has to not only exceed that of seawater, but also be large enough to enable rapid sinking. We thus argue that biofouling cannot be an efficient vertical transport mechanism for small microplastic. However, biofouling of small microplastic may promote the likelihood of its incorporation into sinking marine snow and increase the probability of its ingestion, allowing its transport to depth.

Quantitative review summarizing the effects of oil pollution on subarctic and arctic marine invertebrates.

While meta-analyses are common in the health and some biological sciences, there is a lack of such analyses for petroleum-related marine research. Oil is a highly complex substance consisting of thousands of different compounds. Measurement limitations, different protocols and a lack of standards in recording and reporting various elements of laboratory experiments impede attempts to homogenize and compare data and identify trends. Nevertheless, oil toxicology research would benefit from meta-analyses, through which we could develop meaningful research questions and design robust experiments. Here we report findings from an effort to quantitatively summarize results from oil toxicology studies on arctic and subarctic marine invertebrates. We discovered that the vast majority of studies was conducted on crustaceans, followed by molluscs. Analyzing the sensitivity of response measures across taxa we found that the most sensitive responses tend to rank low in ecological relevance, while less sensitive response measures tend to be more ecologically relevant. We further uncovered that crustaceans appear to be more sensitive to mechanically dispersed than chemically dispersed oil while the opposite seems true for molluscs, albeit not statistically significant. Both crustaceans and molluscs show a higher sensitivity to fresh than to weathered oil. No differences in the sensitivities of crustacean life stages were found. However, due to a lack of data, many questions remain unanswered. Our study revealed that while trends in responses can be elucidated, heterogeneous experimental protocols and reporting regimes prevent a proper meta-analysis.

Planktonic Aggregates as Hotspots for Heterotrophic Diazotrophy: The Plot Thickens.

Biological dinitrogen (N2) fixation is performed solely by specialized bacteria and archaea termed diazotrophs, introducing new reactive nitrogen into aquatic environments. Conventionally, phototrophic cyanobacteria are considered the major diazotrophs in aquatic environments. However, accumulating evidence indicates that diverse non-cyanobacterial diazotrophs (NCDs) inhabit a wide range of aquatic ecosystems, including temperate and polar latitudes, coastal environments and the deep ocean. NCDs are thus suspected to impact global nitrogen cycling decisively, yet their ecological and quantitative importance remain unknown. Here we review recent molecular and biogeochemical evidence demonstrating that pelagic NCDs inhabit and thrive especially on aggregates in diverse aquatic ecosystems. Aggregates are characterized by reduced-oxygen microzones, high C:N ratio (above Redfield) and high availability of labile carbon as compared to the ambient water. We argue that planktonic aggregates are important loci for energetically-expensive N2 fixation by NCDs and propose a conceptual framework for aggregate-associated N2 fixation. Future studies on aggregate-associated diazotrophy, using novel methodological approaches, are encouraged to address the ecological relevance of NCDs for nitrogen cycling in aquatic environments.

Emiliania huxleyi-Bacteria Interactions under Increasing CO2 Concentrations.

The interactions established between marine microbes, namely phytoplankton-bacteria, are key to the balance of organic matter export to depth and recycling in the surface ocean. Still, their role in the response of phytoplankton to rising CO2 concentrations is poorly understood. Here, we show that the response of the cosmopolitan Emiliania huxleyi (E. huxleyi) to increasing CO2 is affected by the coexistence with bacteria. Specifically, decreased growth rate of E. huxleyi at enhanced CO2 concentrations was amplified in the bloom phase (potentially also related to nutrient concentrations) and with the coexistence with Idiomarina abyssalis (I. abyssalis) and Brachybacterium sp. In addition, enhanced CO2 concentrations also affected E. huxleyi's cellular content estimates, increasing organic and decreasing inorganic carbon, in the presence of I. abyssalis, but not Brachybacterium sp. At the same time, the bacterial isolates only survived in coexistence with E. huxleyi, but exclusively I. abyssalis at present CO2 concentrations. Bacterial species or group-specific responses to the projected CO2 rise, together with the concomitant effect on E. huxleyi, might impact the balance between the microbial loop and the export of organic matter, with consequences for atmospheric carbon dioxide.

The Complexity of Spills: The Fate of the Deepwater Horizon Oil.

The Deepwater Horizon oil spill was the largest, longest-lasting, and deepest oil accident to date in US waters. As oil and natural gas jetted from release points at 1,500-m depth in the northern Gulf of Mexico, entrainment of the surrounding ocean water into a buoyant plume, rich in soluble hydrocarbons and dispersed microdroplets of oil, created a deep (1,000-m) intrusion layer. Larger droplets of liquid oil rose to the surface, forming a slick of mostly insoluble, hydrocarbon-type compounds. A variety of physical, chemical, and biological mechanisms helped to transform, remove, and redisperse the oil and gas that was released. Biodegradation removed up to 60% of the oil in the intrusion layer but was less efficient in the surface slick, due to nutrient limitation. Photochemical processes altered up to 50% (by mass) of the floating oil. The surface oil expression changed daily due to wind and currents, whereas the intrusion layer flowed southwestward. A portion of the weathered surface oil stranded along shorelines. Oil from both surface and intrusion layers were deposited onto the seafloor via sinking marine oil snow. The biodegradation rates of stranded or sedimented oil were low, with resuspension and redistribution transiently increasing biodegradation. The subsequent research efforts increased our understanding of the fate of spilled oil immensely, with novel insights focusing on the importance of photooxidation, the microbial communities driving biodegradation, and the formation of marine oil snow that transports oil to the seafloor.

Biological response to dissolved versus dispersed oil.

The water-soluble compounds of oil (e.g. low molecular weight PAHs) dissolve as a function of their physicochemical properties and environmental conditions, while the non-soluble compounds exist as dispersed droplets. Both the chemical and physical form of oil will affect the biological response. We present data from a mesocosm study comparing the microbial response to the water-soluble fraction (WSF), versus a water-accommodated fraction of oil (WAF), which contains both dispersed and dissolved oil components. WAF and WSF contained similar concentrations of low molecular weight PAHs, but concentrations of 4- and 5-ring PAHs were higher in WAF compared to WSF. Microbial communities were significantly different between WSF and WAF treatments, primary productivity was reduced more in WSF than in WAF, and concentrations of transparent exopolymeric particles were highest in WSF and lowest in the controls. These differences highlight the importance of dosing strategy for mesocosm and toxicity tests.

Inter-laboratory calibration of estimated oil equivalent (EOE) concentrations of a water accommodated fraction (WAF) of oil and a chemically enhanced WAF (CEWAF).

Chemical characterization of the presence of oil in environmental samples are performed using methods of varying complexity. Extraction of samples with an organic solvent and analysis by fluorescence spectrometry has been shown to be a rapid and effective screening technique for petroleum in the environment. During experiments, rapid analysis of oil by fluorescence provides the opportunity for researchers to modify the experimental conditions in real time. Estimated Oil Equivalents (EOE) relies on the fluorescence measurement of the aromatic compounds to estimate the oil concentration. The present intercalibration study was designed to investigate whether different fluorometer instruments can reliably measure EOE and whether the results are intercomparable. Additionally, the need for extraction of oil compounds into an organic solvent was investigated. Three different fluorometers were used in three different laboratories: a Horiba Aqualog, a Turner Trilogy and a Shimadzu Spectrofluorophotometer RF-1501. Results from these different instruments showed excellent agreement for EOE determinations. A very high correlation was found between the EOE results obtained with Aqualog Horiba and Turner Trilogy (r2 = 0.9999), with no significant differences between the mean EOE results (t-test, p = 0.30), and the Aqualog Horiba and Shimadzu (r2 = 0.995) fluorometers, with no statistically difference between the EOE results obtained by the two instruments (p = 0.40).

Response of natural phytoplankton communities exposed to crude oil and chemical dispersants during a mesocosm experiment.

During the 2010 Deepwater Horizon oil spill, the chemical dispersant Corexit was applied over vast areas of the Gulf of Mexico. Marine phytoplankton play a key role in aggregate formation through the production of extracellular polymeric materials (EPS), an important step in the biological carbon pump. This study examined the impacts of oil and dispersants on the composition and physiology of natural marine phytoplankton communities from the Gulf of Mexico during a 72-hour mesocosm experiment and consequences to carbon export. The communities were treated using the water accommodated fraction (WAF) of oil, which was produced by adding Macondo surrogate oil to natural seawater and mixed for 24 h in the dark. A chemically enhanced WAF (CEWAF) was made in a similar manner, but using a mixture of oil and the dispersant Corexit in a 20:1 ratio as well as a diluted CEWAF (DCEWAF). Phytoplankton communities exposed to WAF showed no significant changes in PSII quantum yield (Fv/Fm) or electron transfer rates (ETRmax) compared to Control communities. In contrast, both Fv/Fm and ETRmax declined rapidly in communities treated with either CEWAF or DCEWAF. Analysis of other photophysiological parameters showed that photosystem II (PSII) antenna size and PSII connectivity factor were not altered by exposure to DCEWAF, suggesting that processes downstream of PSII were affected. The eukaryote community composition in each experimental tank was characterized at the end of the 72 h exposure time using 18S rRNA sequencing. Diatoms dominated the communities in both the control and WAF treatments (52 and 56% relative abundance respectively), while in CEWAF and DCEWAF treatments were dominated by heterotrophic Euglenozoa (51 and 84% respectively). Diatoms made up the largest relative contribution to the autotrophic eukaryote community in all treatments. EPS concentration was four times higher in CEWAF tanks compared to other treatments. Changes in particle size distributions (a proxy for aggregates) over time indicated that a higher degree of particle aggregation occurred in both the CEWAF and DCEWAF treatments than the WAF or Controls. Our results demonstrate that chemically dispersed oil has more negative impacts on photophysiology, phytoplankton community structure and aggregation dynamics than oil alone, with potential implications for export processes that affect the distribution and turnover of carbon and oil in the water column.

Effects of salinity and transparent exopolymer particles on formation of aquatic aggregates and their association with norovirus.

Human noroviruses (NoVs) are responsible for 50% of food-related disease outbreaks and are notably associated with shellfish consumption. Despite the detrimental health impacts of human NoV-contaminated seafood to public health, there is a lack of knowledge on the physicochemical conditions that govern NoV transmission in aquatic ecosystems. In the present study, we investigated the propensity for NoVs to associate with aquatic aggregates, which have been shown to efficiently deliver nano-sized particles to shellfish. Specific physicochemical conditions characteristic of shellfish cultivation waters, specifically salinity and transparent exopolymer particles (TEP), were targeted in this study for investigating aggregate formation and NoV association dynamics. Murine norovirus (MNV) was used in aggregation experiments as a model surrogate for NoVs. Results demonstrate increased aggregate formation as a function of increasing salinity and TEP concentrations, as well as greater numbers of MNV genomes incorporated into aggregates under conditions that favor aggregation. As aggregate formation was enhanced in waters representing optimal conditions for shellfish production, specifically saline and high TEP waters, the implications to virus transport and shellfish food safety are profound: more aggregates implies increased scavenging of virus particles from surrounding waters and therefor greater risk for bivalve contamination with nano-sized pathogens. These novel data provide insight into where and when NoVs are most likely to be ingested by shellfish via contaminated aggregates, thereby informing best management and water quality monitoring practices aimed at providing safe seafood to consumers.

How the dispersant Corexit impacts the formation of sinking marine oil snow.

The vertical transport of sinking marine oil snow (MOS) and oil-sediment aggregations (OSA) during the Deepwater Horizon (DwH) spill contributed appreciably to the unexpected, and exceptional accumulation of oil on the seafloor. However, the role of the dispersant Corexit in mediating oil-sedimentation is still controversial. Here we demonstrate that the formation of diatom MOS is enhanced by chemically undispersed oil, but inhibited by Corexit-dispersed oil. Nevertheless, the sedimentation rate of oil may at times be enhanced by Corexit application, because of an elevated oil content per aggregate when Corexit is used. A conceptual framework explains the seemingly contradictory effects of Corexit application on the sedimentation of oil and marine particles. The redistribution of oil has central ecological implications, and future decisions on mediating measures or damage assessment will have to take the formation of sinking, oil-laden, marine snow into account.

Elevated pCO2 enhances bacterioplankton removal of organic carbon.

Factors that affect the removal of organic carbon by heterotrophic bacterioplankton can impact the rate and magnitude of organic carbon loss in the ocean through the conversion of a portion of consumed organic carbon to CO2. Through enhanced rates of consumption, surface bacterioplankton communities can also reduce the amount of dissolved organic carbon (DOC) available for export from the surface ocean. The present study investigated the direct effects of elevated pCO2 on bacterioplankton removal of several forms of DOC ranging from glucose to complex phytoplankton exudate and lysate, and naturally occurring DOC. Elevated pCO2 (1000-1500 ppm) enhanced both the rate and magnitude of organic carbon removal by bacterioplankton communities compared to low (pre-industrial and ambient) pCO2 (250 -~400 ppm). The increased removal was largely due to enhanced respiration, rather than enhanced production of bacterioplankton biomass. The results suggest that elevated pCO2 can increase DOC consumption and decrease bacterioplankton growth efficiency, ultimately decreasing the amount of DOC available for vertical export and increasing the production of CO2 in the surface ocean.

Evaluating Carbonate System Algorithms in a Nearshore System: Does Total Alkalinity Matter?

Ocean acidification is a threat to many marine organisms, especially those that use calcium carbonate to form their shells and skeletons. The ability to accurately measure the carbonate system is the first step in characterizing the drivers behind this threat. Due to logistical realities, regular carbonate system sampling is not possible in many nearshore ocean habitats, particularly in remote, difficult-to-access locations. The ability to autonomously measure the carbonate system in situ relieves many of the logistical challenges; however, it is not always possible to measure the two required carbonate parameters autonomously. Observed relationships between sea surface salinity and total alkalinity can frequently provide a second carbonate parameter thus allowing for the calculation of the entire carbonate system. Here, we assessed the rigor of estimating total alkalinity from salinity at a depth <15 m by routinely sampling water from a pier in southern California for several carbonate system parameters. Carbonate system parameters based on measured values were compared with those based on estimated TA values. Total alkalinity was not predictable from salinity or from a combination of salinity and temperature at this site. However, dissolved inorganic carbon and the calcium carbonate saturation state of these nearshore surface waters could both be estimated within on average 5% of measured values using measured pH and salinity-derived or regionally averaged total alkalinity. Thus we find that the autonomous measurement of pH and salinity can be used to monitor trends in coastal changes in DIC and saturation state and be a useful method for high-frequency, long-term monitoring of ocean acidification.

Natural oil slicks fuel surface water microbial activities in the northern Gulf of Mexico.

We conducted a series of roller tank incubations with surface seawater from the Green Canyon oil reservoir, northern Gulf of Mexico, amended with either a natural oil slick (GCS-oil) or pristine oil. The goal was to test whether bacterial activities of natural surface water communities facilitate the formation of oil-rich marine snow (oil snow). Although oil snow did not form during any of our experiments, we found specific bacterial metabolic responses to the addition of GCS-oil that profoundly affected carbon cycling within our 4-days incubations. Peptidase and ß-glucosidase activities indicative of bacterial enzymatic hydrolysis of peptides and carbohydrates, respectively, were suppressed upon the addition of GCS-oil relative to the non-oil treatment, suggesting that ascending oil and gas initially inhibits bacterial metabolism in surface water. Biodegradation of physically dispersed GCS-oil components, indicated by the degradation of lower molecular weight n-alkanes as well as the rapid transformation of particulate oil-carbon (C: N >40) into the DOC pool, led to the production of carbohydrate- and peptide-rich degradation byproducts and bacterial metabolites such as transparent exopolymer particles (TEP). TEP formation was highest at day 4 in the presence of GCS-oil; in contrast, TEP levels in the non-oil treatment already peaked at day 2. Cell-specific enzymatic activities closely followed TEP concentrations in the presence and absence of GCS-oil. These results demonstrate that the formation of oil slicks and activities of oil-degrading bacteria result in a temporal offset of microbial cycling of organic matter, affecting food web interactions and carbon cycling in surface waters over cold seeps.