Egg boons: central components of marine fatty acid food webs.

Food web relationships are traditionally defined in terms of the flow of key elements, such as carbon, nitrogen, and phosphorus, and their role in limiting production. There is growing recognition that availability of important biomolecules, such as fatty acids, may exert controls on secondary production that are not easily explained by traditional element-oriented models. Essential fatty acids (EFAs) are required by most organisms for proper physiological function but are manufactured almost entirely by primary producers. Therefore, the flow of EFAs, especially docosahexaenoic acid (DHA), eicosapentaenoic acid (EPA), and arachidonic acid (ARA), through aquatic food webs is critical for ecosystem functioning. A meta-analysis of data on the EFA content of marine organisms reveals that individual eggs of marine animals have exceptionally high concentrations of EFAs, and that superabundances of eggs released in temporally and spatially discrete patches create rich, but temporary, nutritional resources for egg predators, called "egg boons." Mortality rates of fish eggs are disproportionately higher than animals of similar size, and those eggs are consumed by predators, both larger and smaller than the adults that produce the eggs. Thus, egg boons are a major trophic pathway through which EFAs are repackaged and redistributed, and they are among the few pathways that run counter to the main direction of trophic flow. Egg boons can transport EFAs across ecosystems through advection of patches of eggs and spawning migrations of adults. Recognizing the significance of egg boons to aquatic food webs reveals linkages and feedbacks between organisms and environments that have important implications for understanding how food webs vary in time and space. Examples are given of top-down, bottom-up, and lateral control mechanisms that could significantly alter food webs through their effects on eggs. Our results suggest that trophodynamic food web models should include EFAs generally, and egg production and egg EFA content in particular.

Urea uptake and carbon fixation by marine pelagic bacteria and archaea during the Arctic summer and winter seasons.

How Arctic climate change might translate into alterations of biogeochemical cycles of carbon (C) and nitrogen (N) with respect to inorganic and organic N utilization is not well understood. This study combined 15N uptake rate measurements for ammonium, nitrate, and urea with 15N- and 13C-based DNA stable-isotope probing (SIP). The objective was to identify active bacterial and archeal plankton and their role in N and C uptake during the Arctic summer and winter seasons. We hypothesized that bacteria and archaea would successfully compete for nitrate and urea during the Arctic winter but not during the summer, when phytoplankton dominate the uptake of these nitrogen sources. Samples were collected at a coastal station near Barrow, AK, during August and January. During both seasons, ammonium uptake rates were greater than those for nitrate or urea, and nitrate uptake rates remained lower than those for ammonium or urea. SIP experiments indicated a strong seasonal shift of bacterial and archaeal N utilization from ammonium during the summer to urea during the winter but did not support a similar seasonal pattern of nitrate utilization. Analysis of 16S rRNA gene sequences obtained from each SIP fraction implicated marine group I Crenarchaeota (MGIC) as well as Betaproteobacteria, Firmicutes, SAR11, and SAR324 in N uptake from urea during the winter. Similarly, 13C SIP data suggested dark carbon fixation for MGIC, as well as for several proteobacterial lineages and the Firmicutes. These data are consistent with urea-fueled nitrification by polar archaea and bacteria, which may be advantageous under dark conditions.

Microbial Community Response to Terrestrially Derived Dissolved Organic Matter in the Coastal Arctic.

Warming at nearly twice the global rate, higher than average air temperatures are the new 'normal' for Arctic ecosystems. This rise in temperature has triggered hydrological and geochemical changes that increasingly release carbon-rich water into the coastal ocean via increased riverine discharge, coastal erosion, and the thawing of the semi-permanent permafrost ubiquitous in the region. To determine the biogeochemical impacts of terrestrially derived dissolved organic matter (tDOM) on marine ecosystems we compared the nutrient stocks and bacterial communities present under ice-covered and ice-free conditions, assessed the lability of Arctic tDOM to coastal microbial communities from the Chukchi Sea, and identified bacterial taxa that respond to rapid increases in tDOM. Once thought to be predominantly refractory, we found that ∼7% of dissolved organic carbon and ∼38% of dissolved organic nitrogen from tDOM was bioavailable to receiving marine microbial communities on short 4 - 6 day time scales. The addition of tDOM shifted bacterial community structure toward more copiotrophic taxa and away from more oligotrophic taxa. Although no single order was found to respond universally (positively or negatively) to the tDOM addition, this study identified 20 indicator species as possible sentinels for increased tDOM. These data suggest the true ecological impact of tDOM will be widespread across many bacterial taxa and that shifts in coastal microbial community composition should be anticipated.

How much crude oil can zooplankton ingest? Estimating the quantity of dispersed crude oil defecated by planktonic copepods.

We investigated and quantified defecation rates of crude oil by 3 species of marine planktonic copepods (Temora turbinata, Acartia tonsa, and Parvocalanus crassirostris) and a natural copepod assemblage after exposure to mechanically or chemically dispersed crude oil. Between 88 and 100% of the analyzed fecal pellets from three species of copepods and a natural copepod assemblage exposed for 48 h to physically or chemically dispersed light crude oil contained crude oil droplets. Crude oil droplets inside fecal pellets were smaller (median diameter: 2.4-3.5 µm) than droplets in the physically and chemically dispersed oil emulsions (median diameter: 6.6 and 8.0 µm, respectively). This suggests that copepods can reject large crude oil droplets or that crude oil droplets are broken into smaller oil droplets before or during ingestion. Depending on the species and experimental treatments, crude oil defecation rates ranged from 5.3 to 245 ng-oil copepod(-1) d(-1), which represent a mean weight-specific defecation rate of 0.026 µg-oil µg-Ccopepod(1) d(-1). Considering a dispersed crude oil concentration commonly found in the water column after oil spills (1 µl L(-1)) and copepod abundances in high productive coastal areas, copepods may defecate ∼ 1.3-2.6 mg-oil m(-3) d(-1), which would represent ∼ 0.15%-0.30% of the total dispersed oil per day. Our results indicate that ingestion and subsequent defecation of crude oil by planktonic copepods has a small influence on the overall mass of oil spills in the short term, but may be quantitatively important in the flux of oil from surface water to sediments and in the transfer of low-solubility, toxic petroleum hydrocarbons into food webs after crude oil spills in the sea.

Influence of UVB radiation on the lethal and sublethal toxicity of dispersed crude oil to planktonic copepod nauplii.

Toxic effects of petroleum to marine zooplankton have been generally investigated using dissolved petroleum hydrocarbons and in the absence of sunlight. In this study, we determined the influence of natural ultraviolet B (UVB) radiation on the lethal and sublethal toxicity of dispersed crude oil to naupliar stages of the planktonic copepods Acartia tonsa, Temora turbinata and Pseudodiaptomus pelagicus. Low concentrations of dispersed crude oil (1 µL L(-1)) caused a significant reduction in survival, growth and swimming activity of copepod nauplii after 48 h of exposure. UVB radiation increased toxicity of dispersed crude oil by 1.3-3.8 times, depending on the experiment and measured variables. Ingestion of crude oil droplets may increase photoenhanced toxicity of crude oil to copepod nauplii by enhancing photosensitization. Photoenhanced sublethal toxicity was significantly higher when T. turbinata nauplii were exposed to dispersant-treated oil than crude oil alone, suggesting that chemical dispersion of crude oil may promote photoenhanced toxicity to marine zooplankton. Our results demonstrate that acute exposure to concentrations of dispersed crude oil and dispersant (Corexit 9500) commonly found in the sea after oil spills are highly toxic to copepod nauplii and that natural levels of UVB radiation substantially increase the toxicity of crude oil to these planktonic organisms. Overall, this study emphasizes the importance of considering sunlight in petroleum toxicological studies and models to better estimate the impact of crude oil spills on marine zooplankton.

Novel insight into the role of heterotrophic dinoflagellates in the fate of crude oil in the sea.

Although planktonic protozoans are likely to interact with dispersed crude oil after a spill, protozoan-mediated processes affecting crude oil pollution in the sea are still not well known. Here, we present the first evidence of ingestion and defecation of physically or chemically dispersed crude oil droplets (1-86 µm in diameter) by heterotrophic dinoflagellates, major components of marine planktonic food webs. At a crude oil concentration commonly found after an oil spill (1 µL L(-1)), the heterotrophic dinoflagellates Noctiluca scintillans and Gyrodinium spirale grew and ingested ~0.37 µg-oil µg-C(dino)(-1) d(-1), which could represent ~17% to 100% of dispersed oil in surface waters when heterotrophic dinoflagellates are abundant or bloom. Egestion of faecal pellets containing crude oil by heterotrophic dinoflagellates could contribute to the sinking and flux of toxic petroleum hydrocarbons in coastal waters. Our study indicates that crude oil ingestion by heterotrophic dinoflagellates is a noteworthy route by which petroleum enters marine food webs and a previously overlooked biological process influencing the fate of crude oil in the sea after spills.