Tara L. Connelly

Urea Uptake and Carbon Fixation by Marine Pelagic Bacteria and Archaea during the Arctic Summer and Winter Seasons

ABSTRACT 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.

Effect of temperature on rates of ammonium uptake and nitrification in the western coastal Arctic during winter, spring, and summer

Biogeochemical rate processes in the Arctic are not currently well constrained, and there is very limited information on how rates may change as the region warms. Here we present data on the sensitivity of ammonium (NH4+) uptake and nitrification rates to short-term warming. Samples were collected from the Chukchi Sea off the coast of Barrow, Alaska, during winter, spring, and summer and incubated for 24 h in the dark with additions of 15NH4+ at −1.5, 6, 13, and 20°C. Rates of NH4+ uptake and nitrification were measured in conjunction with bacterial production. In all seasons, NH4+ uptake rates were highest at temperatures similar to current summertime conditions but dropped off with increased warming, indicative of psychrophilic (i.e., cold-loving) microbial communities. In contrast, nitrification rates were less sensitive to temperature and were higher in winter and spring compared to summer. These findings suggest that as the Arctic coastal ecosystem continues to warm, NH4+ assimilation may become increasingly important, relative to nitrification, although the magnitude of NH4+ assimilation would be still be lower than nitrification.

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-Cdino−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.

Elemental composition, total lipid content, and lipid class proportions in zooplankton from the benthic boundary layer of the Beaufort Sea shelf (Canadian Arctic)

Biochemical analyses such as lipid class and elemental composition can inform us about a species’ role in community energetics and nutrient cycling. The accumulation of lipid-rich energy stores affects the elemental composition and stoichiometry of animal tissues, and this relationship is especially relevant to zooplankton at higher latitudes due to increased seasonal lipid storage. However, due to sampling difficulties, the elemental composition and energy storage capabilities of polar, benthic boundary layer zooplankton are poorly known. We determined elemental and lipid class compositions for 26 taxa of benthic boundary layer zooplankton from the Beaufort Sea shelf. Elemental composition as a percentage of dry weight ranged 21–56% for carbon (C), 4–11% for nitrogen (N), and 0.1–1.1% for phosphorus (P) across all taxa. C concentration and C:N were positively correlated with the storage lipids triacylglycerols (TG) and wax esters/steryl esters (WE/SE) and negatively correlated with membrane lipids (phospholipids and sterols). Most taxa had high levels of storage lipids, generally TG. High levels of WE/SE were found in the copepod Calanus hyperboreus (>90% of total lipid) and the chaetognath Eukrohnia hamata (72%). In contrast, the chaetognath Parasagitta elegans had only minor proportions of both TG and WE/SE. The high levels of storage lipids in most taxa indicate that feeding behavior of benthic boundary layer zooplankton on the Beaufort Sea shelf is tightly linked with seasonal pulses of epipelagic production. This is the first report on the biochemical composition of most of the amphipod and mysid taxa presented here.

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.

Fatty-acid biomarkers and tissue-specific turnover: validation from a controlled feeding study in juvenile Atlantic croaker Micropogonias undulatus.

Fatty-acid (FA) profiles of liver and muscle tissue from juvenile Atlantic croaker Micropogonias undulatus were examined over a 15 week diet-switch experiment to establish calibration coefficients (CC) and improve understanding of consumer-diet relationships for field applications. Essential FAs [docosahexaenoic acid (DHA), 22:6n-3 and eicosapentaenoic acid (EPA) , 20:5n-3] decreased and 18:2n-6 increased in tissues of M. undulatus fed diets with increasing proportions of terrestrial v. marine lipid sources. Non-linear models used to estimate the incorporation rate and days to saturation of per cent 18:2n-6 in tissues showed that livers incorporated 18:2n-6 faster than muscle, but the proportions of 18:2n-6 in muscle were higher. CCs were established to determine proportions of FA deposition in tissues relative to diet. Many CCs were consistent amongst diet treatments, despite growth and dietary differences. The CCs can be used to discern FA modification and retention within tissues and as tools for future quantitative estimates of diet histories. Incorporation rates and CCs of 18:2n-6 were applied to a sub-set of field samples of wild M. undulatus to understand habitat use and feeding ecology. Altogether, these results suggest that FAs provide a time-integrated measure of diet in aquatic food webs and are affected by tissue type, growth rate and the influence of mixed diets.

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.