Giant sponge grounds of Central Arctic seamounts are associated with extinct seep life.

The Central Arctic Ocean is one of the most oligotrophic oceans on Earth because of its sea-ice cover and short productive season. Nonetheless, across the peaks of extinct volcanic seamounts of the Langseth Ridge (87°N, 61°E), we observe a surprisingly dense benthic biomass. Bacteriosponges are the most abundant fauna within this community, with a mass of 460 g C m-2 and an estimated carbon demand of around 110 g C m-2 yr-1, despite export fluxes from regional primary productivity only sufficient to provide <1% of this required carbon. Observed sponge distribution, bulk and compound-specific isotope data of fatty acids suggest that the sponge microbiome taps into refractory dissolved and particulate organic matter, including remnants of an extinct seep community. The metabolic profile of bacteriosponge fatty acids and expressed genes indicate that autotrophic symbionts contribute significantly to carbon assimilation. We suggest that this hotspot ecosystem is unique to the Central Arctic and associated with extinct seep biota, once fueled by degassing of the volcanic mounts.

Role of carbonate burial in Blue Carbon budgets.

Calcium carbonates (CaCO3) often accumulate in mangrove and seagrass sediments. As CaCO3 production emits CO2, there is concern that this may partially offset the role of Blue Carbon ecosystems as CO2 sinks through the burial of organic carbon (Corg). A global collection of data on inorganic carbon burial rates (Cinorg, 12% of CaCO3 mass) revealed global rates of 0.8 TgCinorg yr-1 and 15-62 TgCinorg yr-1 in mangrove and seagrass ecosystems, respectively. In seagrass, CaCO3 burial may correspond to an offset of 30% of the net CO2 sequestration. However, a mass balance assessment highlights that the Cinorg burial is mainly supported by inputs from adjacent ecosystems rather than by local calcification, and that Blue Carbon ecosystems are sites of net CaCO3 dissolution. Hence, CaCO3 burial in Blue Carbon ecosystems contribute to seabed elevation and therefore buffers sea-level rise, without undermining their role as CO2 sinks.

Benthic primary producers are key to sustain the Wadden Sea food web: stable carbon isotope analysis at landscape scale.

Coastal food webs can be supported by local benthic or pelagic primary producers and by the import of organic matter. Distinguishing between these energy sources is essential for our understanding of ecosystem functioning. However, the relative contribution of these components to the food web at the landscape scale is often unclear, as many studies lack good taxonomic and spatial resolution across large areas. Here, using stable carbon isotopes, we report on the primary carbon sources for consumers and their spatial variability across one of the world's largest intertidal ecosystems (Dutch Wadden Sea; 1460 km2 intertidal surface area), at an exceptionally high taxonomic (178 species) and spatial resolution (9,165 samples from 839 locations). The absence of overlap in δ13 C values between consumers and terrestrial organic matter suggests that benthic and pelagic producers dominate carbon input into this food web. In combination with the consistent enrichment of benthic primary producers (δ13 C -16.3‰) relative to pelagic primary producers (δ13 C -18.8) across the landscape, this allowed the use of a two-food-source isotope-mixing model. This spatially resolved modelling revealed that benthic primary producers (microphytobenthos) are the most important energy source for the majority of consumers at higher trophic levels (worms, molluscs, crustaceans, fish, and birds), and thus to the whole food web. In addition, we found large spatial heterogeneity in the δ13 C values of benthic primary producers (δ13 C -19.2 to -11.5‰) and primary consumers (δ13 C -25.5 to -9.9‰), emphasizing the need for spatially explicit sampling of benthic and pelagic primary producers in coastal ecosystems. Our findings have important implications for our understanding of the functioning of ecological networks and for the management of coastal ecosystems.

Factors controlling the oral bioaccessibility of anthropogenic Pb in polluted soils.

In human risk assessment, ingestion of soil is considered a major route of toxic Pb exposure. A large body of research has focussed on the measurement of the 'total' Pb contents in sediment, soil and dust as a measure for the exposure to lead. We report that Pb bioaccessibility (i.e. the maximum bioavailability), determined with an in vitro test, does not necessarily depend on the total Pb content. In contrast, the Pb bioaccessibility is initially controlled by the chemical form and particle size of the Pb source, which in turn determine its solubility. Furthermore, when anthropogenic Pb resides within the soil, it may form new, more stable, minerals and/or binds to organic matter, clay, reactive iron or other reactive phases, changing its bioaccessibility. The bioaccessible Pb fraction of 28 soils, polluted with various Pb sources (including residues of Pb bullets and pellets, car battery Pb, city waste and diffuse Pb), was determined with an in vitro-test and varied from 0.5% to 79.0% of total Pb. The highest Pb bioaccessibility (60.7% to 79.0%) was measured in soils polluted with residues of Pb bullets and pellets (shooting range), while the lowest Pb bioaccessibility (0.5%-8.3%) was measured in soils polluted with city waste (including remnants of Pb glazed potsherds and rooftiles, Pb based paint flakes, and Pb sheets). Bioaccessibility of Pb was correlated with pH, organic matter and reactive Fe. These results indicate that soil characteristics play an important role in the oral bioaccessibility of lead in polluted soils. Instead of basing human risk assessment solely on total Pb contents we propose to incorporate in vitro bioaccessibility tests, taking factors such as soil pH, organic matter content and reactive iron content into account. This approach will result in a better insight into the actual risks of Pb polluted soils to children.

Reconstruction of historical atmospheric Pb using Dutch urban lake sediments: a Pb isotope study.

Lake sediments provide a record of atmospheric Pb deposition and changes in Pb isotope composition. To our knowledge, such an approach has not previously been performed in The Netherlands or linked to national air monitoring data. Results are presented for Pb content and isotope composition of (137)Cs dated lake sediments from 2 Dutch urban lakes. Between 1942 and 2002A.D. anthropogenic atmospheric Pb deposition rates in the two lakes varied from 12±2 to 69±16µgcm(-2)year(-1). The rise and fall of leaded gasoline is clearly reflected in the reconstructed atmospheric Pb deposition rates. After the ban on leaded gasoline, late 1970s/early 1980s, atmospheric Pb deposition rates decreased rapidly in the two urban lakes and the relative contributions of other anthropogenic Pb sources - incinerator ash (industrial Pb) and coal/galena - increased sharply. Atmospheric Pb deposition rates inferred from the lake record a clear relationship with nearby measured annual mean air Pb concentrations. Based on this relationship it was estimated that air Pb concentrations between 1942 and 2002A.D. varied between 5 and 293ng/m(3).

The lead (Pb) isotope signature, behaviour and fate of traffic-related lead pollution in roadside soils in The Netherlands.

In this study the origin, behaviour and fate of anthropogenic Pb in sandy roadside soils were assessed by measuring soil characteristics, Pb isotope composition and content. In 1991 and 2003 samples were taken at different depth intervals at approximately 8 and 75 m from two highways in The Netherlands. The Pb isotope composition of the litter layer ((206)Pb/(207)Pb=1.12-1.14) differs from the deeper soil samples ((206)Pb/(207)Pb=1.20-1.21). Based on a mixing model it is concluded that the samples contain two Pb sources: natural Pb and anthropogenic Pb, the latter mainly derived from gasoline. (206)Pb/(207)Pb ratios demonstrate that the roadside soils were polluted to a depth of ~15 cm. Within this depth interval, anthropogenic Pb content is associated with organic matter. Although Pb pollution only reached a depth of ~15 cm, this does not mean that the topsoils retain all anthropogenic Pb. Due to the low pH and negligible binding capacity of soils at depths >15 cm, anthropogenic Pb migrated towards groundwater after reaching depths of >15 cm. The Pb isotope composition of the groundwater ((206)Pb/(207)Pb=1.135-1.185) establishes that groundwater is polluted with anthropogenic Pb. The contribution of anthropogenic Pb to the groundwater varies between ~30 and 100%. Based on the difference in soil Pb content and Pb isotope compositions over a period of 12 years, downward Pb migration is calculated to vary from 72 ± 95 to 324 ± 279 mg m(-2)y(-1). Assuming that the downward Pb flux is constant over time, it is calculated that 35-90% of the atmospherically delivered Pb has migrated to the groundwater.

Carbon conversion and metabolic rate in two marine sponges.

The carbon metabolism of two marine sponges, Haliclona oculata and Dysidea avara, has been studied using a 13C isotope pulse-chase approach. The sponges were fed 13C-labeled diatoms (Skeletonema costatum) for 8 h and they took up between 75 and 85%. At different times, sponges were sampled for total 13C enrichment, and fatty acid (FA) composition and 13C enrichment. Algal biomarkers present in the sponges were highly labeled after feeding but their labeling levels decreased until none was left 10 days after enrichment. The sponge-specific FAs incorporated 13C label already during the first day and the amount of 13C label inside these FAs kept increasing until 3 weeks after labeling. The algal-derived carbon captured by the sponges during the 8-h feeding period was thus partly respired and partly metabolized during the weeks following. Apparently, sponges are able to capture enough food during short periods to sustain longer-term metabolism. The change of carbon metabolic rate of fatty acid synthesis due to mechanical damage of sponge tissue was studied by feeding sponges with 13C isotope-labeled diatom (Pheaodactylum tricornutum) either after or before damaging and tracing back the 13C content in the damaged and healthy tissue. The filtration and respiration in both sponges responded quickly to damage. The rate of respiration in H. oculata reduced immediately after damage, but returned to its initial level after 6 h. The 13C data revealed that H. oculata has a higher metabolic rate in the tips where growth occurs compared to the rest of the tissue and that the metabolic rate is increased after damage of the tissue. For D. avara, no differences were found between damaged and non-damaged tissue. However, the filtration rate decreased directly after damage.

Uncertainties in ecological, chemical and physiological parameters of a bioaccumulation model: implications for internal concentrations and tissue based risk quotients.

Bioaccumulation models predict internal contaminant concentrations (c(i)) using ecological, chemical and physiological parameters. Here we analyse the effect of uncertainties on these parameters on bioaccumulation model predictions. Simultaneously considering the uncertainties on all these parameters in a bioaccumulation model resulted in uncertainty ranges of c(i) that increased with the octanol water partition coefficient K(ow) and reached maxima of up to 1.25 log units for mesozooplankton and up to 1.45 log units fish at logK(ow)=8. A global sensitivity analysis (SA) was performed to rank the contribution of different parameters to the observed uncertainty. The SA demonstrated that this interspecies difference resulted predominantly from uncertain production rates of fish. The K(ow), the water concentration and organic carbon-octanol proportionality constant were important drivers of uncertainty on c(i) for both species. A tissue based risk quotient (RQ(tissue)) combining uncertainty on c(i) with realistic tissue based effect thresholds indicated that fish were up to 10 times more probable to have RQ(tissue)>1 than mesozooplankton, depending on the considered threshold value. Conventional exposure based risk quotients were up to 5 times less probable to exceed one than were corresponding RQ(tissue), and this for both species.

Inferring chemical effects on carbon flows in aquatic food webs: methodology and case study.

The majority of ecotoxicological enclosure experiments monitor species abundances at different chemical concentrations. Here, we present a new modelling approach that estimates changes in food web flows from such data and show that population- and food web level effects are revealed that are not apparent from abundance data alone. For the case of cypermethrin in freshwater enclosures, photosynthesis and excretion (d(-1)) of phytoplankton at 3.643 microg L(-1) cypermethrin were 30% lower and 100% higher than in the control, respectively. The ingestion rate of mesozooplankton (d(-1)) was 6 times higher in the treated enclosures than in the control as food concentration increased with insecticide exposure. With increasing cypermethrin concentrations, nanoflagellates progressively relied on phytoplankton as their main food source, which rendered the food web less stable. We conclude that this tool has excellent potential to analyse the wealth of enclosure data as it only needs species abundance and general constraints.

A versatile method for stable carbon isotope analysis of carbohydrates by high-performance liquid chromatography/isotope ratio mass spectrometry.

We have developed a method to analyze stable carbon isotope ((13)C/(12)C) ratios in a variety of carbohydrates using high-performance liquid chromatography/isotope ratio mass spectrometry (HPLC/IRMS). The chromatography is based on strong anion-exchange columns with low strength NaOH eluents. An eluent concentration of 1 mM resulted in low background signals and good separation of most of the typical plant neutral carbohydrates. We also show that more strongly bound carbohydrates such as acidic carbohydrates can be separated by inclusion of NO(3) (-) as an inorganic pusher ion in the eluent. Analyses of neutral carbohydrate concentrations and their stable carbon isotope ratios are shown for plant materials and marine sediment samples both at natural abundance and for (13)C-enriched samples. The main advantage of HPLC/IRMS analysis over traditional gas chromatography based methods is that no derivatization is needed resulting in simple sample treatment and improved accuracy and reproducibility.

Impacts of atmospheric anthropogenic nitrogen on the open ocean.

Increasing quantities of atmospheric anthropogenic fixed nitrogen entering the open ocean could account for up to about a third of the ocean's external (nonrecycled) nitrogen supply and up to approximately 3% of the annual new marine biological production, approximately 0.3 petagram of carbon per year. This input could account for the production of up to approximately 1.6 teragrams of nitrous oxide (N2O) per year. Although approximately 10% of the ocean's drawdown of atmospheric anthropogenic carbon dioxide may result from this atmospheric nitrogen fertilization, leading to a decrease in radiative forcing, up to about two-thirds of this amount may be offset by the increase in N2O emissions. The effects of increasing atmospheric nitrogen deposition are expected to continue to grow in the future.

Human trampling as short-term disturbance on intertidal mudflats: effects on macrofauna biodiversity and population dynamics of bivalves.

The effect of physical disturbance in the form of trampling on the benthic environment of an intertidal mudflat was investigated. Intense trampling was created as unintended side-effect by benthic ecologists during field experiments in spring and summer 2005, when a mid-shore area of 25 × 25 m was visited twice per month by on average five researchers for a period of 8 months. At the putatively-impacted location (I) (25 × 25 m) and two nearby control locations (Cs) (25 × 25 m each), three sites (4 × 4 m) were randomly selected and at each site, three plots (50 × 50 cm) were sampled after 18 and 40 days from the end of the disturbance. Multivariate and univariate asymmetrical analyses tested for changes in the macrofaunal assemblage, biomass of microphytobenthos and various sediment properties (grain-size, water content, NH4 and NO3 concentrations in the pore water) between the two control locations (Cs) and the putatively-impacted location (I). There were no detectable changes in the sediment properties and microphytobenthos biomass, but variability at small scale was observed. Microphytobenthos and NH4 were correlated at I to the number of footprints, as estimated by the percentage cover of physical depressions. This indicated that trampling could have an impact at small scales, but more investigation is needed. Trampling, instead, clearly modified the abundance and population dynamics of the clam Macoma balthica (L.) and the cockle Cerastoderma edule (L.). There was a negative impact on adults of both species, probably because footsteps directly killed or buried the animals, provoking asphyxia. Conversely, trampling indirectly enhanced recruitment rate of M. balthica, while small-sized C. edule did not react to the trampling. It was likely that small animals could recover more quickly because trampling occurred during the growing season and there was a continuous supply of larvae and juveniles. In addition, trampling might have weakened negative adult-juvenile interactions between adult cockles and juvenile M. balthica, thus facilitating the recruitment. Our findings indicated that human trampling is a relevant source of disturbance for the conservation and management of mudflats. During the growing season recovery can be fast, but in the long-term it might lead towards the dominance of M. balthica to the cost of C. edule, thereby affecting ecosystem functioning.

Stable isotopes and biomarkers in microbial ecology.

The use of biomarkers in combination with stable isotope analysis is a new approach in microbial ecology and a number of papers on a variety of subjects have appeared. We will first discuss the techniques for analysing stable isotopes in biomarkers, primarily gas chromatography-combustion-isotope ratio mass spectrometry, and then describe a number of applications in microbial ecology based on 13C. Natural abundance isotope ratios of biomarkers can be used to study organic matter sources utilised by microorganisms in complex ecosystems and for identifying specific groups of bacteria like methanotrophs. Addition of labelled substrates in combination with biomarker analysis enables direct identification of microbes involved in specific processes and also allows for the incorporation of bacteria into food web studies. We believe that the full potential of the technique in microbial ecology has just started to be exploited.

Spatial distribution and inhibition by ammonium of methane oxidation in intertidal freshwater marshes.

In two intertidal marshes, the vertical distribution in the sediment and inhibition by ammonium of methane oxidation were investigated by slurry incubation experiments. The two sites differ in their dominant vegetation type, i.e., reed and bulrush, and in their heights above sea level. The reed site was elevated with respect to the bulrush site, resulting in a lower frequency and duration of flooding and, consequently, a higher potential for methane oxidation. Methane oxidation decreased with depth in the bulrush and reed slurries, although methane oxidation associated with root material from the bulrush plants increased with depth. Reed root material had a limited capacity for methane oxidation and showed no significant increase with depth. Inhibition of methane oxidation by ammonium was observed in all samples and depended on methane and ammonium concentrations. Increasing ammonium concentrations resulted in greater inhibition, and increasing methane concentrations resulted in less. Ammonium concentrations had to exceed methane concentrations by at least 30-fold to become effective for inhibition. This ratio was found only in the surface layer of the sediment. Hence, the ecological relevance for ammonium inhibition of methane oxidation in intertidal marshes is rather limited and is restricted to the surface layer. Nitrate production was restricted to the 0- to 5-cm-depth slurries.