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Dispersants can aid dispersion and biodegradation of oil in seawater, but the wider ecotoxicological effects of oil and dispersant to the base of marine food webs is unclear. Here we apply a metatranscriptomic approach to identify molecular responses of a natural marine microbial eukaryotic community to oil and chemically dispersed oil. Oil exposure stimulated the upregulation of ketogenesis in the eukaryotic community, which may alleviate carbon- and energy-limitation and reduce oxidative stress. In contrast, a chemically dispersed oil treatment stimulated eukaryotic genes and pathways consistent with nitrogen and oxygen depletion. These results suggest that the addition of dispersant may elevate bacterial biodegradation of crude oil, indirectly increasing competition for nitrogen between prokaryotic and eukaryotic communities as oxygen consumption induces bacterial anaerobic respiration and denitrification. Eukaryotic microbial communities may mitigate some of the negative effects of oil exposure such as reduced photosynthesis and elevated oxidative stress, through ketosis, but the addition of dispersant to the oil fundamentally alters the environmental and ecological conditions and therefore the biochemical response of the eukaryotic community., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Diatoms play a key role in the marine carbon cycle with their high primary productivity and release of exudates such as extracellular polymeric substances (EPS) and transparent exopolymeric particles (TEP). These exudates contribute to aggregates (marine snow) that rapidly transport organic material to the seafloor, potentially capturing contaminants like petroleum components. Ocean acidification (OA) impacts marine organisms, especially those that utilize inorganic carbon for photosynthesis and EPS production. Here we investigated the response of the diatom Thalassiosira pseudonana grown to present day and future ocean conditions in the presence of a water accommodated fraction (WAF and OAWAF) of oil and a diluted chemically enhanced WAF (DCEWAF and OADCEWAF). T. pseudonana responded to WAF/DCEWAF but not OA and no multiplicative effect of the two factors (i.e., OA and oil/dispersant) was observed. T. pseudonana released more colloidal EPS (< 0.7 μm to > 3 kDa) in the presence of WAF/DCEWAF/OAWAF/OADCEWAF than in the corresponding Controls. Colloidal EPS and particulate EPS in the oil/dispersant treatments have higher protein-to-carbohydrate ratios than those in the control treatments, and thus are likely stickier and have a greater potential to form aggregates of marine oil snow. More TEP was produced in response to WAF than in Controls; OA did not influence its production. Polyaromatic hydrocarbon (PAH) concentrations and distributions were significantly impacted by the presence of dispersants but not OA. PAHs especially Phenanthrenes, Anthracenes, Chrysenes, Fluorenes, Fluoranthenes, Pyrenes, Dibenzothiophenes and 1-Methylphenanthrene show major variations in the aggregate and surrounding seawater fraction of oil and oil plus dispersant treatments. Studies like this add to the current knowledge of the combined effects of aggregation, marine snow formation, and the potential impacts of oil spills under ocean acidification scenarios., Competing Interests: The authors have declared that no competing interests exist.

The Deepwater Horizon oil spill released millions of barrels of crude oil into the Gulf of Mexico, and saw widespread use of the chemical dispersant Corexit. We assessed the role of traits, such as cell size, cell wall, motility, and mixotrophy on the growth and photosynthetic response of 15 phytoplankton taxa to oil and Corexit. We collected growth and photosynthetic data on five algal cultures. These responses could be separated into resistant (Tetraselmis astigmatica, Ochromonas sp., Heterocapsa pygmaea) and sensitive (Micromonas pusilla, Prorocentrum minimum). We combined this data with 10 species previously studied and found that cell size is most important in determining the biomass response to oil, whereas motility/mixotrophy is more important in the dispersed oil. Our analysis accounted for a third of the variance observed, so further work is needed to identify other factors that contribute to oil resistance., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020. Published by Elsevier Ltd.)

Species-level variability has made it difficult to determine the relative sensitivity of phytoplankton to oil and mixtures of oil and dispersant. Here we develop a phytoplankton group sensitivity index using ribosome sequence data that we apply to a mesocosm experiment in which a natural microbial community was exposed to oil and two oil-dispersant mixtures. The relative sensitivity of four phytoplankton taxonomic groups, diatoms, dinoflagellates, green algae, and Chrysophytes, was computed using the log of the ratio of the number of species that increase to the number that decrease in relative abundance in the treatment relative to the control. The index indicates that dinoflagellates are the most sensitive group to oil and oil-dispersant treatments while the Chrysophytes benefit under oil exposure compared to the other groups examined. The phytoplankton group sensitivity index can be generally applied to quantify and rank the relative sensitivity of diverse microbial groups to environmental conditions and pollutants., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Here, we report results from a 15-day mesocosm experiment examining changes in estimated oil equivalents (EOEs), n-alkanes (n-C 10 to n-C 35 ), polycyclic aromatic hydrocarbons (PAHs) and petroleum biomarkers. Water accommodated fractions (WAF) of oil and diluted chemically enhanced WAF (DCEWAF) were prepared and concentrations of oil residues determined on day 0, 3 and 15, respectively. Significant removals of n-alkane and PAHs were observed starting from day 3. The n-C 17 /pristane and n-C 18 /phytane ratios suggested that the n-alkane removal was due to biodegradation in the mesocosms. The ratios of C2-dibenzothiophenes/C2-phenanthrenes (D2/P2) and C3-dibenzothiophenes/C3-phenanthrenes (D3/P3) were found to be stable through the experiment. DCEWAF treatment had longer half-lives for most n-alkanes but shorter half-lives for most PAHs than the WAF treatment. Most petroleum biomarkers were stable throughout the experiment. However, depletion of TAS (tricyclic aromatic steroids) was observed on day 15 of DCEWAF treatment., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2019. Published by Elsevier Ltd.)

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., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Large amounts of oil containing mucous-like marine snow formed in surface waters adjacent to the Deepwater Horizon spill that was implicated in oil delivery to the seafloor. However, whether chemical dispersants that were used increased or decreased the oil incorporation and sedimentation efficiency, and how exopolymeric substances (EPS) are involved in this process remains unresolved. To investigate the microbial responses to oil and dispersants in different oceanic settings, indicated by EPS production, petro- and non-petro carbon sedimentation, four mesocosm (M) experiments were conducted: 1) nearshore seawater with a natural microbial consortia (M2); 2) offshore seawater with f/20 nutrients (M3); 3) coastal seawater with f/20 nutrients (M4); 4) nearshore seawater with a natural microbial consortia for a longer duration (M5). Four treatments were conducted in M2, M3 and M4 whereas only three in M5: 1) a water accommodated fraction of oil (WAF), 2) a chemically-enhanced WAF prepared with Corexit (CEWAF, not in M5), 3) a 10-fold diluted CEWAF (DCEWAF); and 4) controls. Overall, oil and dispersants input, nutrient and microbial biomass addition enhanced EPS production. Dispersant addition tended to induce the production of EPS with higher protein/carbohydrate (P/C) ratios, irrespective of oceanic regions. EPS produced in M4 was generally more hydrophobic than that produced in M3. The P/C ratio of EPS in both the aggregate and the colloidal fraction was a key factor that regulated oil contribution to sinking aggregates, based on the close correlation with %petro-carbon in these fractions. In the short term (4-5 days), both the petro and non-petro carbon sedimentation efficiencies showed decreasing trends when oil/dispersants were present. In comparison, in the longer-term (16 days), petro-carbon sedimentation efficiency was less influenced by dispersants, possibly due to biological and physicochemical changes of the components of the oil-EPS-mineral phase system, which cooperatively controlled the sinking velocities of the aggregates., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Spilled oil treated with Corexit dispersant can cause unintended impacts on marine environment systems including altering marine organic matter dynamics; however, impacts on microgels and marine oil snow (MOS) formation are still debated and remain to be fully understood. Extracellular polymeric substances (EPS) are a major source of marine organic carbon for MOS and microgel formation. EPS initial aggregation plays key roles in the oil degrading process and various biogeochemical reactions. Here we used four types of EPS with water accommodated fraction (WAF), chemically-enhanced WAF (CEWAF) and Corexit, to represent potential situations during oil spills and post-application of Corexit. We found that Corexit alone can inhibit EPS aggregation and disperse pre-existing microgels. CEWAF can enhance EPS aggregation with efficiency by up to 80%-100% and more aggregates accumulated within the air-water interface. Additionally, more hydrophobic EPS aggregates showed high resistance to Corexit dispersion while hydrophilic EPS were more sensitive. Effects of oil spills on marine gel particle formation are primarily determined by chemical characteristics (hydrophobicity and protein content) of the constituent EPS. This study offers unique insights for organic particle dynamics and identifies controlling factors for MOS or gel particles associated with oil spills and Corexit dispersant used., (Copyright © 2018 Elsevier B.V. All rights reserved.)

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 (F v /F m ) or electron transfer rates (ETR max ) compared to Control communities. In contrast, both F v /F m and ETR max 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., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Phytoplankton alter their macromolecule composition in response to changing environmental conditions. Often these changes are consistent and can be used as indicators to predict their exposure to a given condition. FTIR-spectroscopy is a powerful tool that provides rapid snapshot of microbial samples. We used FTIR to develop signature macromolecular composition profiles of three cultures: Skeletonema costatum, Emiliania huxleyi, and Navicula sp., exposed to chemically enhanced water accommodated oil fraction (CEWAF) in artificial seawater and control. Using a multivariate model created with a Partial Least Square Discriminant Analysis of the FTIR-spectra, classification of CEWAF exposed versus control samples was possible. This model was validated using aggregate samples from a mesocosm study. Analysis of spectra and PCA-loadings plot showed changes to carbohydrates and proteins in response to CEWAF. Overall we developed a robust multivariate model that can be used to identify if a phytoplankton sample has been exposed to oil with dispersant., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Corexit, an EPA-approved chemical dispersant, was intensively used during the 2010 Deepwater Horizon Oil Spill in the Gulf of Mexico. Current studies surrounding Corexit have mainly focused on its toxicity and oil removal capacity. The potential impact of Corexit on the surface ocean carbon dynamics has remained largely unknown. The spontaneous assembly of DOM (dissolved organic matter) polymers into microgels (POM, particulate organic matter) was demonstrated previously that it can influence various critical processes, such as colloidal pump, microbial loops, and nutrition availability in the surface ocean. Here, we report that Corexit alone can significantly inhibit DOM microgel formation and reduce the stability of pre-existing microgels. However, Corexit and oil, Chemically Enhanced Water Accommodated Fraction (CEWAF), could effectively facilitate DOM microgel formation. The unanticipated disturbance of Corexit and oil spills on the critical DOM-POM continuum warrant particular caution and thus should be considered for future application of Corexit during oil spills., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Human alterations to the marine environment such as an oil spill can induce oxidative stress in phytoplankton. Exposure to oil has been shown to be lethal to most phytoplankton species, but some are able to survive and grow at unaffected or reduced growth rates, which appears to be independent of the class and phylum of the phytoplankton and their ability to consume components of oil heterotrophically. The goal of this article is to test the role of core metabolism plasticity in the oil‐resisting ability of phytoplankton. Experiments were performed on the oil‐ resistant chlorophyte, Dunaliella tertiolecta, in control and water accommodated fractions of oil, with and without metabolic inhibitors targeting the core metabolic pathways. We observed that inhibiting pathways such as photosynthetic electron transport (PET) and pentose‐phosphate pathway were lethal; however, inhibition of pathways such as mitochondrial electron transport and cyclic electron transport caused growth to be arrested. Pathways such as photorespiration and Kreb's cycle appear to play a critical role in the oil‐tolerating ability of D. tertiolecta. Analysis of photo‐physiology revealed reduced PET under inhibition of photorespiration but not Kreb's cycle. Further studies showed enhanced flux through Kreb's cycle suggesting increased energy production and photorespiration counteract oxidative stress. Lastly, reduced extracellular carbohydrate secretion under oil exposure indicated carbon and energy conservation, which together with enhanced flux through Kreb's cycle played a major role in the survival of D. tertiolecta under oil exposure by meeting the additional energy demands. Overall, we present data that suggest the role of phenotypic plasticity of multiple core metabolic pathways in accounting for the oxidative stress tolerating ability of certain phytoplankton species. [ABSTRACT FROM AUTHOR]

Biological oil weathering facilitated by specialized heterotrophic microbial communities plays a key role in the fate of petroleum hydrocarbon in the ocean. The most common methods of assessing oil biodegradation involve (i) measuring changes in the composition and concentration of oil over time and/or (ii) biological incubations with stable or radio-labelled substrates. Both methods provide robust and invaluable information on hydrocarbon biodegradation pathways; however, they also require extensive sample processing and are expensive in nature. More convenient ways to assess activities within microbial oil degradation networks involve measuring extracellular enzyme activity. This perspective article synthesizes previously published results from studies conducted in the aftermath of the 2010 Deepwater Horizon (DwH) oil spill in the northern Gulf of Mexico (nGoM), to test the hypothesis that fluorescence assays of esterases, including lipase activity, are sensitive indicators for microbial oil degradation in the ocean. In agreement with the rates and patterns of enzyme activity in oil-contaminated seawater and sediments in the nGoM, we found close correlations between esterase activity measured by means of methylumbeliferyl (MUF) oleate and MUF butyrate hydrolysis, and the concentration of petroleum hydrocarbons in two separate laboratory incubations using surface (<1 m) and deep nGoM waters (>1200 m). Correlations between esterase activities and oil were driven by the presence of chemical dispersants, suggesting a connection to the degree of oil dissolution in the medium. Our results clearly show that esterase activities measured with fluorogenic substrate proxies are a good indicator for oil biodegradation in the ocean; however, there are certain factors as discussed in this study that need to be taken into consideration while utilizing this approach. [ABSTRACT FROM AUTHOR]

Phytoplankton play a central role in our ecosystems, they are responsible for nearly 50 percent of the global primary productivity and major drivers of macro-elemental cycles in the ocean. Phytoplankton are constantly subjected to stressors, some natural such as nutrient limitation and some manmade such as oil spills. With increasing oil exploration activities in coastal zones in the Gulf of Mexico and elsewhere, an oil spill during nutrient-limited conditions for phytoplankton growth is highly likely. We performed a multifactorial study exposing the diatom Phaeodactylum tricornutum (UTEX 646) to oil and/or dispersants under nitrogen and silica limitation as well as co-limitation of both nutrients. Our study found that treatments with nitrogen limitation (-N and–N-Si) showed overall lower growth and chlorophyll a, lower photosynthetic antennae size, lower maximum photosynthetic efficiency, lower protein in exopolymeric substance (EPS), but higher connectivity between photosystems compared to non-nitrogen limited treatments (-Si and +N+Si) in almost all the conditions with oil and/or dispersants. However, certain combinations of nutrient limitation and oil and/or dispersant differed from this trend indicating strong interactive effects. When analyzed for significant interactive effects, the–N treatment impact on cellular growth in oil and oil plus dispersant conditions; and oil and oil plus dispersant conditions on cellular growth in–N-Si and–N treatments were found to be significant. Overall, we demonstrate that nitrogen limitation can affect the oil resistant trait of P. tricornutum, and oil with and without dispersants can have interactive effects with nutrient limitation on this diatom. [ABSTRACT FROM AUTHOR]

The 2010 Deepwater Horizon oil-spill exposed the microbes of Gulf of Mexico to unprecedented amount of oil. Conclusive evidence of the underlying molecular mechanism(s) on the negative effects of oil exposure on certain phytoplankton species such as Thalassiosira pseudonana is still lacking, curtailing our understanding of how oil spills alter community composition. We performed experiments on model diatom T. pseudonana to understand the mechanisms underpinning observed reduced growth and photosynthesis rates during oil exposure. Results show severe impairment to processes upstream of photosynthesis, such as light absorption, with proteins associated with the light harvesting complex damaged while the pigments were unaffected. Proteins associated with photosynthetic electron transport were also damaged, severely affecting photosynthetic apparatus and depriving cells of energy and carbon for growth. Negative growth effects were alleviated when an organic carbon source was provided. Further investigation through proteomics combined with pathway enrichment analysis confirmed the above findings, while highlighting other negatively affected processes such as those associated with ferroxidase complex, high-affinity iron-permease complex, and multiple transmembrane transport. We also show that oxidative stress is not the primary route of negative effects, rather secondary. Overall, this study provides a mechanistic understanding of the cellular damage that occurs during oil exposure to T. pseudonana. [ABSTRACT FROM AUTHOR]

Microbial interactions influence nearly one-half of the global biogeochemical flux of major elements of the marine ecosystem. Despite their ecological importance, microbial interactions remain poorly understood and even less is known regarding the effects of anthropogenic perturbations on these microbial interactions. The Deepwater Horizon oil spill exposed the Gulf of Mexico to ∼4.9 million barrels of crude oil over 87 days. We determined the effects of oil exposure on microbial interactions using short- and long-term microcosm experiments with and without Macondo surrogate oil. Microbial activity determined using radiotracers revealed that oil exposure negatively affected substrate uptake by prokaryotes within 8 h and by eukaryotes over 72 h. Eukaryotic uptake of heterotrophic exopolymeric substances (EPS) was more severely affected than prokaryotic uptake of phototrophic EPS. In addition, our long-term exposure study showed severe effects on photosynthetic activity. Lastly, changes in microbial relative abundances and fewer co-occurrences among microbial species were mostly driven by photosynthetic activity, treatment (control vs. oil), and prokaryotic heterotrophic metabolism. Overall, oil exposure affected microbial co-occurrence and/or interactions possibly by direct reduction in abundance of one of the interacting community members and/or indirect by reduction in metabolism (substrate uptake or photosynthesis) of interacting members. [ABSTRACT FROM AUTHOR]

Concerns on the timing and processes associated with petroleum degradation were raised after the use of Corexit during the Deepwater Horizon oil spill. There is a lack of understanding of the removal of oil associated with flocculate materials to the sediment. Mesocosm studies employing coastal and open-ocean seawater from the Gulf of Mexico were undertaken to examine changes in oil concentration and composition with time. The water accommodated fractions (WAF) and chemically enhanced WAF (CEWAF) produced using Macondo surrogate oil and Corexit were followed over 3–4 days in controlled environmental conditions. Environmental half-lives of estimated oil equivalents (EOE), polycyclic aromatic hydrocarbons (PAH), n-alkanes (C10-C35), isoprenoids pristane and phytane, and total petroleum hydrocarbons (TPH) were determined. EOE and PAH concentrations decreased exponentially following first-order decay rate kinetics. WAF, CEWAF and DCEWAF (a 10X CEWAF dilution) treatments half-lives ranged from 0.9 to 3.2 days for EOE and 0.5 to 3.3 days for PAH, agreeing with estimates from previous mesocosm and field studies. The aliphatic half-lives for CEWAF and DECWAF treatments ranged from 0.8 to 2.0 days, but no half-life for WAF could be calculated as concentrations were below the detection limits. Biodegradation occurred in all treatments based on the temporal decrease of the nC17/pristane and nC18/phytane ratios. The heterogeneity observed in all treatments was likely due to the hydrophobicity of oil and weathering processes occurring at different rates and times. The presence of dispersant did not dramatically change the half-lives of oil. Comparing degradation of oil alone as well as with dispersant present is critical to determine the fate and transport of these materials in the ocean. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Marine snow formation and vertical transport are naturally occurring processes that carry organic matter from the surface to deeper waters, providing food and sequestering carbon. During the Deepwater Horizon well blowout, oil was incorporated with marine snow aggregates, triggering a Marine Oil Snow (MOS) Sedimentation and Flocculent Accumulation (MOSSFA) event, that transferred a significant percentage of the total released oil to the seafloor. An improved understanding of processes controlling MOS formation and MOSSFA events is necessary for evaluating their impacts on the fate of spilled oil. Numerical models and predictive tools capable of providing scientific support for oil spill planning, response, and Natural Resource Damage Assessment are being developed to provide information for weighing the ecological trade-offs of response options. Here we offer considerations for oil spill response and recovery when assessing the potential for a MOSSFA event and provide tools to enhance decision-making. • Sinking MOS can transport oil to the seafloor during and after an oil spill. • MOSSFA should be considered in oil spill response planning, recovery, and NRDA. • Satellite imagery and oil trajectory modeling can help predict MOSSFA events. • Decision tree can be used by oil response community to consider MOSSFA. [ABSTRACT FROM AUTHOR]

The Deepwater Horizon oil spill of 2010 brought the ecology and health of the Gulf of Mexico to the forefront of the public's and scientific community's attention. Not only did we need a better understanding of how this oil spill impacted the Gulf of Mexico ecosystem, but we also needed to apply this knowledge to help assess impacts from perturbations in the region and guide future response actions. Phytoplankton represent the base of the food web in oceanic systems. As such, alterations of the phytoplankton community propagate to upper trophic levels. This review brings together new insights into the influence of oil and dispersant on phytoplankton. We bring together laboratory, mesocosm and field experiments, including insights into novel observations of harmful algal bloom (HAB) forming species and zooplankton as well as bacteria-phytoplankton interactions. We finish by addressing knowledge gaps and highlighting key topics for research in novel areas. [ABSTRACT FROM AUTHOR]

Marine snow was implicated in the transport of oil to the seafloor during the Deepwater Horizon oil spill, but the exact processes remain controversial. In this study, we investigated the concentrations and distributions of the 16 USEPA priority polycyclic aromatic hydrocarbons (PAHs) in marine snow aggregates collected during a mesocosm experiment. Seawater only, oil in a water accommodated fraction (WAF), and Corexit-enhanced WAF (DCEWAF) were incubated for 16 d. Both WAF and DCEWAF aggregates were enriched in heavy molecular weight PAHs but depleted in naphthalene. DCEWAF aggregates had 2.6 times more total 16 PAHs than the WAF (20.5 vs. 7.8 µg/g). Aggregates in the WAF and DCEWAF incorporated 4.4% and 19.3%, respectively of the total PAHs in the mesocosm tanks. Our results revealed that marine snow sorbed and scavenged heavy molecular weight PAHs in the water column and the application of Corexit enhanced the incorporation of PAHs into the sinking aggregates. [ABSTRACT FROM AUTHOR]

Abstract Large amounts of mucous-rich oil-containing marine snow formed in surface waters adjacent to the Deepwater Horizon spill. This marine oil snow (MOS) was implicated in oil delivery to the seafloor. Whether the use of chemical dispersants increased or decreased MOS sedimentation remains controversial. We conducted mesocosm experiments testing the effects of oil and oil plus a dispersant on MOS formation and sedimentation, in coastal seawater. The four treatments used were a surrogate Macondo oil water accommodated fraction ("WAF"), two concentrations of chemically-enhanced WAF ("CEWAF" and diluted CEWAF, "DCEWAF") containing a dispersant and oil, and Controls (no additions). Based on radiocarbon and 13C NMR results, the presence of dispersants enhanced the amounts of petro-carbon being incorporated into the sinking oil-carrying aggregates (aka MOS). However, most of the chemically-dispersed oil preferentially partitioned into the colloidal and suspended particulate fractions rather than into the rapidly forming MOS. Thus the oil and non-petro-carbon sedimentation efficiency in treatments with a dispersant was much lower, compared to those in the Control and WAF treatments, during the four-day mesocosm experiment. Formation of MOS and its subsequent sinking sequestered the oil in two stages: first via terrestrial-derived detritus containing humic compounds, and subsequently via freshly produced material, such as exopolymeric substances produced by phytoplankton and bacteria. Graphical abstract Unlabelled Image Highlights • Δ14C and NMR molecular mixing model to study petrocarbon incorporation to MOS • Corexit enhanced amounts of petro-carbon incorporated into the sinking MOS • Terrestrial-derived detritus played a role in MOS formation and sedimentation • Exopolymeric substances produced by algae and bacteria also influenced this process • Corexit retarded petro- and non-petrocarbon sedimentation in the 4-day mesocosm [ABSTRACT FROM AUTHOR]

Extracellular polymeric substances (EPS) released from marine microbial cells is a major response to stress. Most previous studies focus on the quantity of EPS released from microbes; however, the critical characteristics of EPS composition receive much less attention. Considering the complex nature and diversity of microbes in the marine system also impart distinct behaviors and reactions when exposed to stressful conditions, we cultured four phytoplankton and five bacterial species, to investigate the changes of the chemical composition of EPS under stress from a water-accommodated fraction (WAF) of oil or a chemically-enhanced WAF (CEWAF). Remaining viable cells (DNA amount) of nine marine microbes showed a higher linear negative correlation with increasing protein to carbohydrate (P/C) ratio under CEWAF than WAF treatments. The data also show that EPS with higher protein ratios was released under higher cellular stress levels induced by CEWAF, suggesting that higher protein ratio EPS, which is more hydrophobic, is secreted to physically or chemically ameliorate the hazardous agents. Our findings provide evidence that microbes can actively modify their EPS release and composition in responses to various adverse stress. Collectively, the results support the hypothesis that higher stress not only can trigger more EPS release, but also induce changes in its composition (P/C), thereby affecting environmental processes such as marine oil snow formation and the characteristic of marine organic matter. Unlabelled Image • Oil and Corexit can induce the release of microbial extracellular polymeric substances (EPS). • Higher protein to carbohydrate (P/C) ratio of EPS was released at higher concentrations of oil and Corexit. • Negative correlation between microbial viability and P/C ratio was found under oil and Corexit stress. • A rapid small-volume assay protocol was established to provide assessment for P/C ratio of released EPS. [ABSTRACT FROM AUTHOR]

Extracellular polymeric substances (EPS), produced by microorganisms, are implicated for greatly influencing the fate of environmental contaminants, including oil. Transparent exopolymeric particles (TEP) are gel-like acidic polysaccharide particles that can be stained with Alcian blue, whereas Coomassie stainable particles (CSP) contain proteins and are stained with Coomassie brilliant blue. Marine microgels are reversibly formed from EPS. These terms are often used interchangeably, but they have rarely been measured simultaneously. Mesocosm and bottle experiments provided an opportunity to compare EPS, TEP, CSP and microgels in a water-accommodated fraction (WAF) of oil and seawater (control). Our results reveal that the biopolymers making up EPS, TEP and CSP consisted primarily of polysaccharides and proteins, mostly likely as proteoglycans and glycoproteins. Significant correlations were found between concentrations of TEP-C vs particulate organic carbon (POC), TEP-C vs particulate organic nitrogen (PON), TEP vs EPS, TEP vs CSP, TEP vs carbohydrates, proteins, CSP and carbohydrates, CSP vs proteins, and carbohydrates vs proteins. Chemical analysis of whole particles and colloids yielded both protein and polysaccharides concentrations higher than those in EDTA extraction, thus providing an upper limit of actual EPS contents in the particulate phase. The EPS that was electrostatically held onto particle surfaces (extractable by 1% EDTA) accounted for a minor (~4%) yet relatively constant proportion of TEP. Overall, the concentrations of the three terms ranked in the order of [gels] > [TEP] > [particulate EPS] in the water. Lastly, spectrophotometric methods have limitations in identifying complex or refractory polysaccharides, as evidenced by the comparison between NMR-quantified EPS and the total EPS determined by spectrophotometric methods. This study is the first time these terms were compared in the same sample. They provide useful information when reviewing historical TEP, CSP, EPS data collected field- and laboratory-studies, and provide linkages between them. In addition, they also demonstrate that they could provide complementary information relevant to ecosystem and flux studies. Unlabelled Image • EPS, TEP, CSP and microgels are operational measurements of exopolymeric materials • EPS, TEP, CSP and POC correlated well to each other in mesocosm experiments • EPS measured in the 1% EDTA extractable fraction asseses ~4% of TEP • [gels] > [TEP] > particulate EPS in bottle experiments with Gulf of Mexico waters • Spectrophotometric methods underestimate polysaccharide fraction compared to NMR [ABSTRACT FROM AUTHOR]