Modeling Microplastic Transport in the Marine Environment: Testing Empirical Models of Particle Terminal Sinking Velocity for Irregularly Shaped Particles.

Microplastic (mP) pollution has been indicated as an area of concern in the marine environment. However, there is no consensus on their potential to cause significant ecological harm, and a comprehensive risk assessment of mP pollution is unattainable due to gaps in our understanding of their transport, uptake, and exchange processes. This research considers drag models that have been proposed to calculate the terminal settling velocity of regularly and irregularly shaped particles to assess their applicability in a mP modeling context. The evaluation indicates three models that predict the settling velocity of mPs to a high precision and suggests that an explicit model is the most appropriate for implementation in a mP transport model. This research demonstrates that the mP settling velocity does not vary significantly over time and depth relevant to the scale of an ocean model and that the terminal settling velocity is independent of the initial particle velocity. These findings contribute toward efforts to simulate the vertical transport of mPs in the ocean, which will improve our understanding of the residence time of mPs in the water column and subsequently their availability for uptake into the marine ecosystem.

Chronic Environmental Perturbation Influences Microbial Community Assembly Patterns.

Acute environmental perturbations are reported to induce deterministic microbial community assembly, while it is hypothesized that chronic perturbations promote development of alternative stable states. Such acute or chronic perturbations strongly impact on the pre-adaptation capacity to the perturbation. To determine the importance of the level of microbial pre-adaptation and the community assembly processes following acute or chronic perturbations in the context of hydrocarbon contamination, a model system of pristine and polluted (hydrocarbon-contaminated) sediments was incubated in the absence or presence (discrete or repeated) of hydrocarbon amendment. The community structure of the pristine sediments changed significantly following acute perturbation, with selection of different phylotypes not initially detectable. Conversely, historically polluted sediments maintained the initial community structure, and the historical legacy effect of chronic pollution likely facilitated community stability. An alternative stable state was also reached in the pristine sediments following chronic perturbation, further demonstrating the existence of a legacy effect. Finally, ecosystem functional resilience was demonstrated through occurrence of hydrocarbon degradation by different communities in the tested sites, but the legacy effect of perturbation also strongly influenced the biotic response. This study therefore demonstrates the importance of perturbation chronicity on microbial community assembly processes and reveals ecosystem functional resilience following environmental perturbation.

Characterisation of microbial communities of drill cuttings piles from offshore oil and gas installations.

Drill cuttings (DC) are produced during hydrocarbon drilling operations and are composed of subsurface rock coated with hydrocarbons and drilling fluids. Historic disposal of DC at sea has resulted in the formation of large piles on the seabed that may be left in situ following infrastructure decommissioning. This study provides a first insight into the microbial abundance, diversity and community structure of two DC piles from North Sea oil and gas installations. The abundance of both bacteria and archaea was lower in DC than in surrounding natural sediments. Microbial diversity and richness within DC were low but increased with distance from the piles. Microbial community structure was significantly different in DC piles compared to nearby natural sediments. DC bacterial communities were dominated by Halomonas, Dietzia and Dethiobacter. The presence of such organisms suggests a potential function of hydrocarbon degradation ability and may play an active role in DC pile remediation.

Pressure and temperature effects on deep-sea hydrocarbon-degrading microbial communities in subarctic sediments.

The Hatton-Rockall Basin (North-East Atlantic) is an area with potential for deep-sea (2,900 m) hydrocarbon exploration. Following the Deepwater Horizon oil spill, many investigations into the responses of sediment microbial communities to oil pollution have been undertaken. However, hydrostatic pressure is a parameter that is often omitted due to the technical difficulties associated with conducting experiments at high pressure (>10 MPa). In this study, sediments from 2,900 m in the Hatton-Rockall Basin, following a one-week decompression period in a temperature-controlled room at 5°C, were incubated in factorial combinations of 0.1 and 30 MPa, 5 and 20°C, and contamination with a hydrocarbon mixture or uncontaminated controls to evaluate the effect of these environmental variables on the bacterial community composition. Our results revealed varying effects of pressure, temperature, and oil contamination on the composition of the bacterial community within the sediment. Temperature was the strongest determinant of differences in the bacterial community structure between samples followed by pressure. Oil contamination did not exert a strong change in the sediment bacterial community structure when pressure and temperature conditions were held at in situ levels (30 MPa and 5°C). The γ-proteobacteria Pseudomonas and Colwellia, and several Bacteroidetes dominated communities at 30 MPa. In contrast, hydrocarbon degraders such as Halomonas, Alcanivorax, and Marinobacter decreased in relative abundance at the same pressure. This study highlights the importance of considering hydrostatic pressure in ex situ investigations into hydrocarbon-degrading deepwater microbial communities.

Advective pore-water transport of hydrocarbons in North East Scotland coastal sands.

Oil spills in the marine environment can cause ecosystem-level impacts. Dispersant application as an oil spill response measure leads to the widespread distribution of hydrocarbons in the water column and marine sediments. The North Sea is an area of intense hydrocarbon production and is at risk of oil spills, which are of concern to its benthic ecosystem due to its sediments' high permeability. Here, entrainment of hydrocarbons via pore-water advection into permeable North Sea sands and the associated effect of Superdispersant-25, a commercial oil dispersant, were evaluated in a laboratory. Centrally stirred chambers that induce advective pore-water fluxes in sediments were filled with sediment, seawater and mixtures of oil and Superdispersant-25. Dispersant application had contrasting effects on hydrocarbon interactions with sediment: (1) it reduced accumulation of hydrocarbons in surface sediments and (2) facilitated the entrainment of hydrocarbons up to 8 cm deep into sediments by increasing hydrocarbon solubility in seawater and its subsequent washout or pumping into sediment by pore-water movement. Results here show that dispersant application can have counter-intuitive effects on hydrocarbon interactions with marine sediments and highlight the need for further research in this area to make better informed decision in an oil spill response scenario.

Effect of spatial origin and hydrocarbon composition on bacterial consortia community structure and hydrocarbon biodegradation rates.

Oil reserves in deep-sea sediments are currently subject to intense exploration, with associated risks of oil spills. Previous research suggests that microbial communities from deep-sea sediment (>1000m) can degrade hydrocarbons (HCs), but have a lower degradation ability than shallow (<200m) communities, probably due to in situ temperature. This study aimed to assess the effect of marine origin on microbial HC degradation potential while separating the influence of temperature, and to characterise associated HC-degrading bacterial communities. Microbial communities from 135 and 1000 m deep sediments were selectively enriched on crude oil at in situ temperatures and both consortia were subsequently incubated for 42 days at 20°C with two HC mixtures: diesel fuel or model oil. Significant HC biodegradation occurred rapidly in the presence of both consortia, especially of low molecular weight HCs and was concomitant with microbial community changes. Further, oil degradation was higher with the shallow consortium than with the deep one. Dominant HC-degrading bacteria differed based on both spatial origin of the consortia and supplemented HC types. This study provides evidence for influence of sediment spatial origin and HC composition on the selection and activity of marine HC-degrading bacterial communities and is relevant for future bioremediationdevelopments.

Current status of deepwater oil spill modelling in the Faroe-Shetland Channel, Northeast Atlantic, and future challenges.

As oil reserves in established basins become depleted, exploration and production moves towards relatively unexploited areas, such as deep waters off the continental shelf. The Faroe-Shetland Channel (FSC, NE Atlantic) and adjacent areas have been subject to increased focus by the oil industry. In addition to extreme depths, metocean conditions in this region characterise an environment with high waves and strong winds, strong currents, complex circulation patterns, sharp density gradients, and large small- and mesoscale variability. These conditions pose operational challenges to oil spill response and question the suitability of current oil spill modelling frameworks (oil spill models and their forcing data) to adequately simulate the behaviour of a potential oil spill in the area. This article reviews the state of knowledge relevant to deepwater oil spill modelling for the FSC area and identifies knowledge gaps and research priorities. Our analysis should be relevant to other areas of complex oceanography.

Benthic macroinfaunal community structure, resource utilisation and trophic relationships in two Canadian Arctic Archipelago polynyas.

Climate change driven alterations to patterns of Arctic marine primary production, with increasing phytoplankton- and decreasing ice algal production, have the potential to change the resource utilisation and trophic structure of the benthic communities relying on the algae for food. To predict the benthic responses to dietary changes, we studied the macroinfaunal community compositions, and used the faunal δ13C and δ15N signatures to investigate their main food sources and trophic positions in North Water (NOW) and Lancaster Sound (LS) polynyas in the Canadian Arctic Archipelago. Macroinfaunal density (10 952 ind. m-2) and biomass (3190 mg C m-2) recorded in NOW were higher than previously found in the Arctic at depths >500m, and significantly higher than in LS (8355 ind. m-2 and 2110 mg C m-2). This was attributed to higher particulate organic matter fluxes to seafloor in NOW. Polychaetes were significant taxa at both sites in terms of density and biomass, and in addition crustacean density in NOW and bivalve density in LS were high. Facultative filter and surface deposit feeders were highly prevalent at both sites, suggesting feeding plasticity is a successful strategy for accessing different food sources. The macrofaunal δ13C signatures reflected the signatures of pelagic particulate organic matter at the sites, and an isotope mixing model confirmed phytoplankton as the main food source for most taxa and feeding guilds. The food web length in LS was longer than in NOW (3.2 vs. 2.8 trophic levels). This was attributed to a larger reliance on reworked organic matter by the benthic community in LS, whereas the high export fluxes at the highly productive NOW resulted in higher rates of selective consumption of fresh algal matter. Despite studies suggesting that loss of ice algae from consumer diets in the Arctic might have a negative impact on the benthos, this study suggests that Arctic macrobenthic communities thrive using phytoplankton as their main food source and should thus be able to cope or even benefit from predicted changes to patterns of primary production.

The Variable Influence of Dispersant on Degradation of Oil Hydrocarbons in Subarctic Deep-Sea Sediments at Low Temperatures (0-5 °C).

The microbial degradation of petroleum hydrocarbons at low temperatures was investigated in subarctic deep-sea sediments in the Faroe Shetland Channel (FSC). The effect of the marine oil dispersant, Superdispersant 25 on hydrocarbon degradation was also examined. Sediments collected at 500 and 1000 m depth were spiked with a model oil containing 20 hydrocarbons and incubated at ambient temperature (5 and 0 °C, respectively) with and without marine dispersant. Treatment of sediments with hydrocarbons resulted in the enrichment of Gammaproteobacteria, and specifically the genera Pseudoalteromonas, Pseudomonas, Halomonas, and Cobetia. Hydrocarbon degradation was faster at 5 °C (500 m) with 65-89% of each component degraded after 50 days compared to 0-47% degradation at 0 °C (1000 m), where the aromatic hydrocarbons fluoranthene, anthracene, and Dibenzothiophene showed no degradation. Dispersant significantly increased the rate of degradation at 1000 m, but had no effect at 500 m. There was no statistically significant effect of Superdispersant 25 on the bacterial community structure at either station. These results show that the indigenous bacterial community in the FSC has the capacity to mitigate some of the effects of a potential oil spill, however, the effect of dispersant is ambiguous and further research is needed to understand the implications of its use.

Carbon and Nitrogen Uptake of Calcareous Benthic Foraminifera along a Depth-Related Oxygen Gradient in the OMZ of the Arabian Sea.

Foraminifera are an important faunal element of the benthos in oxygen-depleted settings such as Oxygen Minimum Zones (OMZs) where they can play a relevant role in the processing of phytodetritus. We investigated the uptake of phytodetritus (labeled with (13)C and (15)N) by calcareous foraminifera in the 0-1 cm sediment horizon under different oxygen concentrations within the OMZ in the eastern Arabian Sea. The in situ tracer experiments were carried out along a depth transect on the Indian margin over a period of 4 to 10 days. The uptake of phytodetrital carbon within 4 days by all investigated species shows that phytodetritus is a relevant food source for foraminifera in OMZ sediments. The decrease of total carbon uptake from 540 to 1100 m suggests a higher demand for carbon by species in the low-oxygen core region of the OMZ or less food competition with macrofauna. Especially Uvigerinids showed high uptake of phytodetrital carbon at the lowest oxygenated site. Variation in the ratio of phytodetrital carbon to nitrogen between species and sites indicates that foraminiferal carbon and nitrogen use can be decoupled and different nutritional demands are found between species. Lower ratio of phytodetrital carbon and nitrogen at 540 m could hint for greater demand or storage of food-based nitrogen, ingestion, or hosting of bacteria under almost anoxic conditions. Shifts in the foraminiferal assemblage structure (controlled by oxygen or food availability) and in the presence of other benthic organisms are likely to account for observed changes in the processing of phytodetritus in the different OMZ habitats. Foraminifera dominate the short-term processing of phytodetritus in the OMZ core but are less important in the lower OMZ boundary region of the Indian margin as biological interactions and species distribution of foraminifera change with depth and oxygen levels.

Occurrence of Priming in the Degradation of Lignocellulose in Marine Sediments.

More than 50% of terrestrially-derived organic carbon (terrOC) flux from the continents to the ocean is remineralised in the coastal zone despite its perceived high refractivity. The efficient degradation of terrOC in the marine environment could be fuelled by labile marine-derived material, a phenomenon known as "priming effect", but experimental data to confirm this mechanism are lacking. We tested this hypothesis by treating coastal sediments with 13C-lignocellulose, as a proxy for terrOC, with and without addition of unlabelled diatom detritus that served as the priming inducer. The occurrence of priming was assessed by the difference in lignocellulose mineralisation between diatom-amended treatments and controls in aerobic sediment slurries. Priming of lignocellulose degradation was observed only at the initial stages of the experiment (day 7) and coincided with overall high microbial activity as exemplified by total CO2 production. Lignocellulose mineralisation did not differ consistently between diatom treatments and control for the remaining experimental time (days 14-28). Based on this pattern, we hypothesize that the faster initiation of lignocellulose mineralisation in diatom-amended treatments is attributed to the decomposition of accessible polysaccharide components within the lignocellulose complex by activated diatom degraders. The fact that diatom-degraders contributed to lignocellulose degradation was also supported by the different patterns in 13C-enrichment of phospholipid fatty acids between treatments. Although we did not observe differences between treatments in the total quantity of respired lignocellulose at the end of the experiment, differences in timing could be important in natural ecosystems where the amount of time that a certain compound is subject to aerobic degradation before burial to deeper anoxic sediments may be limited.

How can we identify and communicate the ecological value of deep-sea ecosystem services?

Submarine canyons are considered biodiversity hotspots which have been identified for their important roles in connecting the deep sea with shallower waters. To date, a huge gap exists between the high importance that scientists associate with deep-sea ecosystem services and the communication of this knowledge to decision makers and to the wider public, who remain largely ignorant of the importance of these services. The connectivity and complexity of marine ecosystems makes knowledge transfer very challenging, and new communication tools are necessary to increase understanding of ecological values beyond the science community. We show how the Ecosystem Principles Approach, a method that explains the importance of ocean processes via easily understandable ecological principles, might overcome this challenge for deep-sea ecosystem services. Scientists were asked to help develop a list of clear and concise ecosystem principles for the functioning of submarine canyons through a Delphi process to facilitate future transfers of ecological knowledge. These ecosystem principles describe ecosystem processes, link such processes to ecosystem services, and provide spatial and temporal information on the connectivity between deep and shallow waters. They also elucidate unique characteristics of submarine canyons. Our Ecosystem Principles Approach was successful in integrating ecological information into the ecosystem services assessment process. It therefore has a high potential to be the next step towards a wider implementation of ecological values in marine planning. We believe that successful communication of ecological knowledge is the key to a wider public support for ocean conservation, and that this endeavour has to be driven by scientists in their own interest as major deep-sea stakeholders.

Feeding preferences of abyssal macrofauna inferred from in situ pulse chase experiments.

Climatic fluctuations may significantly alter the taxonomic and biochemical composition of phytoplankton blooms and subsequently phytodetritus, the food source for the majority of deep-sea communities. To examine the response of abyssal benthic communities to different food resources we simulated a food sedimentation event containing diatoms and coccolithophorids at Station M in the NE Pacific. In one set of experiments we measured incorporation of diatomC and coccoN into the macrofauna using isotopically enriched (13)C-diatoms and (15)N-coccolithophores. In a second experiment we measured incorporation of C and N from dual-labelled ((13)C and (15)N) diatoms. The second experiment was repeated 2 months later to assess the effect of seasonality. The simulated food pulses represented additions of 650 - 800 mg C m(-2) and 120 mg N m(-2) to the seafloor. In all cases rapid incorporation of tracer was observed within 4 days, with between 20% and 52% of the macrofauna displaying evidence of enrichment. However, incorporation levels of both diatomC and coccoN were low (<0.05% and 0.005% of the added C and N). Incorporation of labelled diatoms was similar during both June and September suggesting that the community was not food limited during either period. We found no evidence for selective ingestion of the different food types in the metazoan fauna suggesting that macrofauna do not have strong preferences for diatom vs. coccolithophore dominated phytodetrital pulses. C∶N ratios from both experiments suggest that the metazoan macrofauna community appear to have higher C demands and/or assimilation efficiencies compared to N. Concomitantly, the foraminifera preferentially selected for diatomN over coccoN, and we suggest that this may be related to foraminiferal requirements for intracellular nitrate. These experiments provide evidence that abyssal faunal feeding strategies are in part driven by an organism's internal stoichiometric budgets and biochemical requirements.

Negative priming effect on organic matter mineralisation in NE Atlantic slope sediments.

The priming effect (PE) is a complex phenomenon which describes a modification (acceleration or retardation) in the mineralisation rate of refractory organic matter (OM) following inputs of labile material. PEs are well-studied in terrestrial ecosystems owing to their potential importance in the evolution of soil carbon stocks but have been largely ignored in aquatic systems despite the fact that the prerequisite for their occurrence, i.e. the co-existence of labile and refractory OM, is also true for sediments. We conducted stable isotope tracer experiments in continental margin sediments from the NE Atlantic (550-950 m) to study PE occurrence and intensity in relation to labile OM input. Sediment slurries were treated with increasing quantities of the (13)C-labelled diatom Thalassiosira rotula and PE was quantified after 7, 14 and 21 days. There was a stepwise effect of diatom quantity on its mineralisation although mineralisation efficiency dropped with increasing substrate amounts. The addition of diatomaceous OM yielded a negative PE (i.e. retardation of existing sediment OM mineralisation) at the end of the experiment regardless of diatom quantity. Negative PE is often the result of preferential utilisation of the newly deposited labile material by the microbial community ("preferential substrate utilization", PSU) which is usually observed at excessive substrate additions. The fact that PSU and the associated negative PE occurred even at low substrate levels in this study could be attributed to limited amounts of OM subject to priming in our study area (~0.2% organic carbon [OC]) which seems to be an exception among continental slopes (typically >0.5%OC). We postulate that PEs will normally be positive in continental slope sediments and that their intensity will be a direct function of sediment OC content. More experiments with varying supply of substrate targeting C-poor vs. C-rich sediments are needed to confirm these hypotheses.

Macrofauna regulate heterotrophic bacterial carbon and nitrogen incorporation in low-oxygen sediments.

Oxygen minimum zones (OMZs) currently impinge upon >1 million km(2) of sea floor and are predicted to expand with climate change. We investigated how changes in oxygen availability, macrofaunal biomass and retention of labile organic matter (OM) regulate heterotrophic bacterial C and N incorporation in the sediments of the OMZ-impacted Indian continental margin (540-1100 m; [O(2)]=0.35-15 µmol l(-1)). In situ pulse-chase experiments traced (13)C:(15)N-labelled phytodetritus into bulk sediment OM and hydrolysable amino acids, including the bacterial biomarker D-alanine. Where oxygen availability was lowest ([O(2)]=0.35 µmol l(-1)), metazoan macrofauna were absent and bacteria assimilated 30-90% of the labelled phytodetritus within the sediment. At higher oxygen levels ([O(2)]=2-15 µmol l(-1)) the macrofaunal presence and lower phytodetritus retention with the sediment occur concomitantly, and bacterial phytodetrital incorporation was reduced and retarded. Bacterial C and N incorporation exhibited a significant negative relationship with macrofaunal biomass across the OMZ. We hypothesise that fauna-bacterial interactions significantly influence OM recycling in low-oxygen sediments and need to be considered when assessing the consequences of global change on biogeochemical cycles.

Resource quality affects carbon cycling in deep-sea sediments.

Deep-sea sediments cover ~70% of Earth's surface and represent the largest interface between the biological and geological cycles of carbon. Diatoms and zooplankton faecal pellets naturally transport organic material from the upper ocean down to the deep seabed, but how these qualitatively different substrates affect the fate of carbon in this permanently cold environment remains unknown. We added equal quantities of (13)C-labelled diatoms and faecal pellets to a cold water (-0.7 °C) sediment community retrieved from 1080 m in the Faroe-Shetland Channel, Northeast Atlantic, and quantified carbon mineralization and uptake by the resident bacteria and macrofauna over a 6-day period. High-quality, diatom-derived carbon was mineralized >300% faster than that from low-quality faecal pellets, demonstrating that qualitative differences in organic matter drive major changes in the residence time of carbon at the deep seabed. Benthic bacteria dominated biological carbon processing in our experiments, yet showed no evidence of resource quality-limited growth; they displayed lower growth efficiencies when respiring diatoms. These effects were consistent in contrasting months. We contend that respiration and growth in the resident sediment microbial communities were substrate and temperature limited, respectively. Our study has important implications for how future changes in the biochemical makeup of exported organic matter will affect the balance between mineralization and sequestration of organic carbon in the largest ecosystem on Earth.