ALBERT

Description: Fucoidan, a sulfated polysaccharide in brown seaweed, has various biological activities including anti-tumor activity. We investigated the effects of fucoidan on the apoptosis of human promyeloid leukemic cells and fucoidan-mediated signaling pathways. Fucoidan induced apoptosis of HL-60, NB4, and THP-1 cells, but not K562 cells. Fucoidan treatment of HL-60 cells induced activation of caspases-8, -9, and -3, the cleavage of Bid, and changed mitochondrial membrane permeability. Fucoidan-induced apoptosis, cleavage of procaspases, and changes in the mitochondrial membrane permeability were efficiently blocked by depletion of mitogen-activated protein kinase (MAPK) kinase kinase 1 (MEKK1), and inhibitors of MAPK kinase 1 (MEK1) and c Jun NH2-terminal kinase (JNK). The phosphorylation of extracellular signal-regulated kinase 1/2 (ERK1/2) and JNK was increased in fucoidan-treated HL-60, NB4, and THP-1 cells, but not K562 cells. ERK1/2 activation occurred at earlier times than JNK activation and JNK activation was blocked by MEK1 inhibitor. In addition, fucoidan-induced apoptosis was inhibited by addition of glutathione and/or L-NAME, and fucoidan decreased intracellular glutathione concentrations and stimulated nitric oxide (NO) production. Buthionine-[R,S]-sulfoximine rendered HL-60 cells more sensitive to fucoidan. Depletion of MEKK1 and inhibition of MEK1 restored the intracellular glutathione content and abrogated NO production, whereas inhibition of JNK activation by SP600125 restored intracellular glutathione content but failed to inhibit NO production in fucoidan-treated HL-60 cells. These results suggest that activation of MEKK1, MEK1, ERK1/2, and JNK, depletion of glutathione, and production of NO are important mediators in fucoidan-induced apoptosis of human leukemic cells

Description: Mutations or environmental factors that result in reversal of conspicuous left–right asymmetries provide an opportunity to study developmental mechanisms. They may also provide insight into evolutionary changes in asymmetry states within and between species. King crabs (family Lithodidae) have a larger right claw and females typically exhibit a dextrally offset abdomen. Nevertheless, I observed a high incidence of left handedness in laboratory reared box crabs (Lopholithodes foraminatus) and captured the first known egg-bearing female lithodid to exhibit reversed asymmetry. This provided a unique opportunity to characterize the reversed phenotype and to compare the incidence of reversed asymmetry in the offspring of normal and reversed females. Asymmetry of the chelae became apparent in the first postzoeal stage (glaucothoe) and handedness was maintained through subsequent instars. Females with larger left claws developed reversed abdominal asymmetry by the fourth crab stage. No reversed asymmetry was observed in the mandibles of zoea larvae or juveniles of either handedness. The incidence of reversed asymmetry in glaucothoe reared from one reversed and three normal females was high (between 20% and 30%), and independent of maternity (P=0.67). Removal of the right cheliped of fourth stage zoeae, and the major cheliped of glaucothoe, did not reverse the direction of asymmetry. Elevated larval rearing temperature also did not affect the frequency of reversed individuals. This lack of evidence for either heritability or induction of handedness is enigmatic. Further investigation of reversed asymmetry in lithodid crabs may provide valuable insights into the development and evolution of bilateral asymmetries.

Description: Here we show the use of the 210Pb-226Ra excess method to determine the growth rate of two corals from the world's largest known cold-water coral reef, Røst Reef, north of the Arctic circle off Norway. Colonies of each of the two species that build the reef, Lophelia pertusa and Madrepora oculata, were collected alive at 350 m depth using a submersible. Pb and Ra isotopes were measured along the major growth axis of both specimens using low level alpha and gamma spectrometry and trace element compositions were studied. 210Pb and 226Ra differ in the way they are incorporated into coral skeletons. Hence, to assess growth rates, we considered the exponential decrease of initially incorporated 210Pb, as well as the increase in 210Pb from the decay of 226Ra and contamination with 210Pb associated with Mn-Fe coatings that we were unable to remove completely from the oldest parts of the skeletons. 226Ra activity was similar in both coral species, so, assuming constant uptake of 210Pb through time, we used the 210Pb-226Ra chronology to calculate growth rates. The 45.5 cm long branch of M. oculata was 31 yr with an average linear growth rate of 14.4 ± 1.1 mm yr−1 (2.6 polyps per year). Despite cleaning, a correction for Mn-Fe oxide contamination was required for the oldest part of the colony; this correction corroborated our radiocarbon date of 40 yr and a mean growth rate of 2 polyps yr−1. This rate is similar to the one obtained in aquarium experiments under optimal growth conditions.

Description: Transfer of the small gaseous molecules CO2 and NH3 across biological membranes, long thought to occur solely by simple diffusion, is now known to be facilitated by members of two multigene families: aquaporins (AQP) and rhesus (Rh) proteins. Although it is accepted that AQP1 and Rh proteins are involved in CO2 and NH3 transfer, respectively, the idea that a single channel can exhibit selectivity for both gases is controversial. Indeed, studies using the same in vitro model (human red blood cells) have provided evidence both for and against a role for Rh proteins as CO2 channels. Thus, this study was initiated to provide in vivo evidence for a dual function of Rh proteins as ammonia and CO2 channels. Here, we show that in zebrafish (Danio rerio), direct ammonia–CO2 competition experiments in adults or translational knockdowns of Rh proteins in larvae affects both ammonia and CO2 excretion. These results suggest that Rh proteins in zebrafish may be common pathways for transport of ammonia and CO2.

Description: The evolution of internal structure plays a pivotal role in the macroscopic response of granular materials to applied loads. A case in point is the so‐called ‘stress–dilatancy relation’, a cornerstone of Soil Mechanics. Numerous attempts have been made to unravel the connection between stress–dilatancy and the evolution of fabric and contact forces in a deforming granular medium. We re‐examine this connection in light of the recent findings on force chain evolution, in particular, that of collective force chain failure by buckling. This study is focussed on two‐dimensional deformation of dense granular assemblies. Analysis of individual and collective force chain bucklings is undertaken using data from a discrete element simulation. It is shown that the kinematics of force chain buckling lead to significant levels of local dilatation being developed in the buckling force chain particles and their confining first‐ring neighbors. Findings from the simulation are used to guide the development of a lattice model of collective, localized force chain buckling. Consideration of the statics and kinematics of this process yields a new stress–dilatancy relation. The physics of buckling, even at its simplest form, introduces a richness into the stress–dilatancy formulation in a way that preserves the essential aspects of fabric evolution, specifically the buckling mode.

Description: Cormorants, Phalacocorax spp., have great potential to affect recreational fisheries and have increased substantially in abundance over the last decades in UK inland waters. Fabricated refuges provide a potential means of reducing fish losses, yet the benefits of such structures may be marginal if natural shelters are abundant and favoured, or if strong density-dependence limits refuge use. This study examined the efficacy of artificial shelters in mesocosm enclosures that allowed standardised and replicated observations of roach, Rutilus rutilus (L.), distribution. When given a choice between occupying open water, simulated reedbeds and artificial brushwood shelters, roach used brushwood shelters extensively across a range of fish densities. When fish had a choice of occupying open water or reedbeds offering no overhead cover, they actively avoided reedbeds and used open water almost exclusively. Occupation of reedbeds was positively related to the amount of overhead cover they provided. When artificial brushwood shelters and reedbeds offered complete overhead cover, brushwood shelters were occupied twice as much as reedbeds. Artificial shelters may therefore have wide application in stillwaters with abundant reedbeds unless the reedbeds are coupled with extensive overhead cover.

Description: 1. The hypothesis that cyanobacteria have higher optimum growth temperatures and higher growth rates at the optimum as compared to chlorophytes was tested by running a controlled experiment with eight cyanobacteria species and eight chlorophyte species at six different temperatures (20–35 °C) and by performing a literature survey. 2. In the experiment, all organisms except the chlorophyte Monoraphidium minutum grew well up to 35 °C. The chlorophyte Chlamydomonas reinhardtii was the fastest-growing organism over the entire temperature range (20–35 °C). 3. Mean optimum growth temperatures were similar for cyanobacteria (29.2 °C) and chlorophytes (29.2 °C). These results are concordant with published data, yielding slightly higher mean optimum growth temperatures for cyanobacteria (27.2 °C) than for chlorophytes (26.3 °C). 4. Mean growth rates of cyanobacteria at 20 °C (0.42 day−1) were significantly lower than those of chlorophytes at 20 °C (0.62 day−1). However, at all other temperatures, there were no differences between mean growth rates of cyanobacteria and chlorophytes. 5. Mean growth rates at the optimum temperature were similar for cyanobacteria (0.92 day−1) and chlorophytes (0.96 day−1). However, analysis of published data revealed that growth rates of cyanobacteria (0.65 day−1) were significantly lower than those of chlorophytes (0.93 day−1) at their optimum temperatures. 6. Although climate warming will probably lead to an intensification of cyanobacterial blooms, our results indicate that this might not be as a result of higher growth rates of cyanobacteria compared with their chlorophyte competitors. The competitive advantage of cyanobacteria can more likely be attributed to their ability to migrate vertically and prevent sedimentation in warmer and more strongly stratified waters and to their resistance to grazing, especially when warming reduces zooplankton body size.

Description: The Swedish UNESCO World Cultural Heritage site of the Birka and Hovgården Iron Age settlements is well suited for the testing of high-resolution archaeological prospection methods. In May 2006 ground-penetrating radar (GPR) and magnetometer test measurements were conducted at Birka, resulting in data of outstanding quality and new archaeological discoveries, but also demonstrating the need for increased spatial sampling regarding GPR prospection at complex Scandinavian sites. Therefore Birka was selected as a testing ground for a pilot study investigating the suitability of the novel multichannel GPR array system MIRA (MALÅ Imaging Radar Array) for efficient, large-scale GPR surveys with very dense spatial sampling. The study was conducted in May 2008 by MALÅ Geoscience AB in collaboration with the archaeological prospection unit of the Swedish National Heritage Board. The very high-resolution three-dimensional GPR pilot survey demonstrated that it is possible to survey 1 ha and more per day with 8 cm cross-line spacing, mapping archaeological structures in unprecedented resolution, such as postholes of only 25 cm diameter. This paper describes the tested technology and methodology as well as the fieldwork and the results of the study.

Description: In many geotechnical systems, it is not uncommon to observe failure in zones of high localized strain called shear bands. The existing models predict the existence and the extent of these localizations, but provide little insight into the micromechanics within the shear bands. This research captures and compares the variation in microstructure both inside and outside of shear bands that formed in physical laboratory plane strain and companion numerical two-dimensional discrete element method (DEM) biaxial compression experiments. Unsheared and sheared laboratory specimens of Ottawa 20–30 sand of varying dilatancy were solidified using a two-stage resin impregnation procedure. The solidified specimens were sectioned and the resulting surfaces were prepared for microstructure observation using optical bright-field microscopy and stereological analysis. Statistical properties of microstructural parameters for sub-regions in a grid pattern and along predefined inclined zones were determined. Similar measurements were performed on 2D DEM simulation specimens at varying strain levels to characterize the evolution of microstructure with increasing strain. The results showed how differences evolved in the mean, standard deviation, and entropy of void distributions with increasing global strain levels. The results indicate how disorder increases and that the material within the shear band does not adhere to the classical concept of critical state, but reaches a terminal void ratio that is largely a function of initial void ratio. Furthermore, there appears to be a transition zone between the far field and the fully formed shear block, as opposed to an abrupt delineation as is traditionally inferred.

Description: Core HU97048-007PC was recovered from the continental Labrador Sea slope at a water depth of 945 m, 250km seaward from the mouth of Cumberland Sound, and 400km north of Hudson Strait. Cumberland Sound is a structural trough partly floored by Cretaceous mudstones and Paleozoic carbonates. The record extends from 10 to 58 ka. On-board logging revealed a complex series of lithofacies, including buff-colored detrital carbonate-rich sediments [Heinrich (H)-events] frequently bracketed by black facies. We investigate the provenance of these facies using quantitative X-ray diffraction on drill-core samples from Paleozoic and Cretaceous bedrock from the SE Baffin Island Shelf, and on the〈2-mm sediment fraction in a transect of five cores from Cumberland Sound to the NW Labrador Sea. A sediment unmixing program was used to discriminate between sediment sources, which included dolomite-rich sediments from Baffin Bay, calcite-rich sediments from Hudson Strait and discrete sources from Cumberland Sound. Results indicated that the bulk of the sediment was derived from Cumberland Sound, but Baffin Bay contributed to sediments coeval with H-0 (Younger Dryas), whereas Hudson Strait was the source during H-events 1–4. Contributions from the Cretaceous outcrops within Cumberland Sound bracket H-events, thus both leading and lagging Hudson Strait-sourced H-events.

Description: The history of climate modeling begins with conceptual models, followed in the 19th century by mathematical models of energy balance and radiative transfer, as well as simple analog models. Since the 1950s, the principal tools of climate science have been computer simulation models of the global general circulation. From the 1990s to the present, a trend toward increasingly comprehensive coupled models of the entire climate system has dominated the field. Climate model evaluation and intercomparison is changing modeling into a more standardized, modular process, presenting the potential for unifying research and operational aspects of climate science

Description: Gas hydrate-related bottom-simulating reflectors mark the phase boundary between hydrate and free gas in the subsurface, and therefore may be used to estimate geothermal gradients and hence heat flow. The depth and temperature of the phase boundary depend on the composition of the hydrateforming gas and of the pore fluid. In the absence of direct sampling, these compositions remain unknown. We develop an alternative approach that is less sensitive to compositional uncertainties and can be applied when the bottom-simulating reflector is densely sampled in a region with significant seabed relief. We apply this approach to a three-dimensional seismic dataset from the eastern Black Sea.

Description: Summary 1. Parasitized females in mammals, fish and birds can enhance the immune defence of their offspring by transferring specific antibodies for the embryo. Likewise, social insect mothers transfer immunity despite the fact that invertebrates lack antibodies. 2.  Female trans-generational immune priming is consistent with parental investment theory, because mothers invest more into rearing their offspring than fathers. However, when immune priming is not directly linked to parental care, as is often the case in insects that abandon their eggs after oviposition, both sexes might benefit from protecting their offspring. 3. Using the red flour beetle, Tribolium castaneum, we show that after parental exposure to heat-killed bacteria, trans-generational immune priming occurs through fathers as well as mothers. 4. This novel finding challenges the traditional view that males provide only genes to their offspring in species without paternal care, and raises the possibility of a division of tasks with respect to immune protection between parents.

Description: Here we make an initial step toward the development of an ocean assimilation system that can constrain the modelled Atlantic Meridional Overturning Circulation (AMOC) to support climate predictions. A detailed comparison is presented of 1° and 1/4° resolution global model simulations with and without sequential data assimilation, to the observations and transport estimates from the RAPID mooring array across 26.5° N in the Atlantic. Comparisons of modelled water properties with the observations from the merged RAPID boundary arrays demonstrate the ability of in situ data assimilation to accurately constrain the east-west density gradient between these mooring arrays. However, the presence of an unconstrained "western boundary wedge" between Abaco Island and the RAPID mooring site WB2 (16 km offshore) leads to the intensification of an erroneous southwards flow in this region when in situ data are assimilated. The result is an overly intense southward upper mid-ocean transport (0–1100 m) as compared to the estimates derived from the RAPID array. Correction of upper layer zonal density gradients is found to compensate mostly for a weak subtropical gyre circulation in the free model run (i.e. with no assimilation). Despite the important changes to the density structure and transports in the upper layer imposed by the assimilation, very little change is found in the amplitude and sub-seasonal variability of the AMOC. This shows that assimilation of upper layer density information projects mainly on the gyre circulation with little effect on the AMOC at 26° N due to the absence of corrections to density gradients below 2000 m (the maximum depth of Argo). The sensitivity to initial conditions was explored through two additional experiments using a climatological initial condition. These experiments showed that the weak bias in gyre intensity in the control simulation (without data assimilation) develops over a period of about 6 months, but does so independently from the overturning, with no change to the AMOC. However, differences in the properties and volume transport of North Atlantic Deep Water (NADW) persisted throughout the 3 year simulations resulting in a difference of 3 Sv in AMOC intensity. The persistence of these dense water anomalies and their influence on the AMOC is promising for the development of decadal forecasting capabilities. The results suggest that the deeper waters must be accurately reproduced in order to constrain the AMOC.

Description: Since April 2004 the RAPID array has made continuous measurements of the Atlantic Meridional Overturning Circulation (AMOC) at 26° N. Two key components of this system are Ekman transport zonally integrated across 26° N and western boundary current transport in the Florida Straits. Whilst measurements of the AMOC as a whole are somewhat in their infancy, this study investigates what useful information can be extracted on the variability of the Ekman and Florida Straits transports using the decadal timeseries already available. Analysis is also presented for Sverdrup transports zonally integrated across 26° N. The seasonal cycles of Florida Straits, Ekman and Sverdrup transports are quantified at 26° N using harmonic analysis of annual and semi-annual constituents. Whilst Sverdrup transport shows clear semi-annual periodicity, calculations of seasonal Florida Straits and Ekman transports show substantial interannual variability due to contamination by variability at non-seasonal frequencies; the mean seasonal cycle for these transports only emerges from decadal length observations. The Florida Straits and Ekman mean seasonal cycles project on the AMOC with a combined peak-to-peak seasonal range of 3.5 Sv. The combined seasonal range for heat transport is 0.40 PW. The Florida Straits seasonal cycle possesses a smooth annual periodicity in contrast with previous studies suggesting a more asymmetric structure. No clear evidence is found to support significant changes in the Florida Straits seasonal cycle at sub-decadal periods. Whilst evidence of wind driven Florida Straits transport variability is seen at sub-seasonal and annual periods, a model run from the 1/4° eddy-permitting ocean model NEMO is used to identify an important contribution from internal oceanic variability at sub-annual and interannual periods. The Ekman transport seasonal cycle possesses less symmetric structure, due in part to different seasonal transport regimes east and west of 50 to 60° W. Around 60% of non-seasonal Ekman transport variability occurs in phase section-wide at 26° N and is related to the NAO, whilst Sverdrup transport variability is more difficult to decompose.

Description: During the last decade, global surface temperatures did not increase as rapidly as in the preceding decades. Although relatively small compared to the observed centennial scale global warming, it has renewed interest in understanding and even predicting climate on time scales of decades, and sparked a community initiative on near‐term prediction that will feature in the fifth intergovernmental panel on climate change assessment report. Decadal prediction, however, is in its infancy, with only a few publications existing. This article has three aims. The first is to make the case for decadal prediction. Decadal fluctuations in global climate similar to that of recent decades were observed during the past century. Associated with large regional changes in precipitation and climate extremes, they are of socioeconomic importance. Climate models, which capture some aspects of observed decadal variability, indicate that such variations might be partly predictable. The second aim is to describe the major challenges to skilful decadal climate prediction. One is poor understanding of mechanisms of decadal climate variability, with climate models showing little agreement. Sparse observations in the past, particularly in the ocean, are also a limiting factor to developing and testing of initialization and prediction systems. The third aim is to stress that despite promising initial results, decadal prediction is in a highly experimental stage, and care is needed in interpreting results and utilizing data from such experiments. In the long‐term, decadal prediction has the potential to improve models, reduce uncertainties in climate change projections, and be of socioeconomic benefit. Copyright © 2010 John Wiley & Sons, Ltd.

Description: Recent seismic evidence suggested that most oceanic plate hydration is associated with trench-outer rise faulting prior to subduction. Hydration at trenches may have a significant impact on the subduction zone water cycle. Previous seismic experiments conducted to the northwest of Nicoya Peninsula, Northern Costa Rica, have shown that the subducting Cocos lithosphere is pervasively altered, which was interpreted to be due to both hydration (serpentinization) and fracturing of the crustal and upper-mantle rocks. New seismic wide-angle reflection and refraction data were collected along two profiles, running parallel to the Middle American trench axis offshore of central Nicaragua, revealing lateral changes of the seismic properties of the subducting lithosphere. Seismic structure along both profiles is characterized by low velocities both in the crust and upper mantle. Velocities in the uppermost mantle are found to be in the range 7.3–7.5 km s−1; thus are 8–10 per cent lower than velocities typical for unaltered peridotites and hence confirm the assumption that serpentinization is a common process at the trench-outer rise area offshore of Nicaragua. In addition, a prominent velocity anomaly occurred within the crust beneath two seamounts. Here, velocity reduction may indicate increased porosity and perhaps permeability, supporting the idea that seamounts serve as sites for water percolation and circulation.

Description: The rugged submarine topography of the Azores supports a diverse heterozoan association resulting in intense biotically-controlled carbonate-production and accumulation. In order to characterise this cold-water (C) factory a 2-year experiment was carried out in the southern Faial Channel to study the biodiversity of hardground communities and for budgeting carbonate production and degradation along a bathymetrical transect from the intertidal to bathyal 500 m depth. Seasonal temperatures peak in September (above a thermocline) and bottom in March (stratification diminishes) with a decrease in amplitude and absolute values with depth, and tidal-driven short-term fluctuations. Measured seawater stable isotope ratios and levels of dissolved nutrients decrease with depth, as do the calcium carbonate saturation states. The photosynthetic active radiation shows a base of the euphotic zone in ~70 m and a dysphotic limit in ~150 m depth. Bioerosion, being primarily a function of light availability for phototrophic endoliths and grazers feeding upon them, is ~10 times stronger on the illuminated upside versus the shaded underside of substrates in the photic zone, with maximum rates in the intertidal (−631 g/m2/yr). Rates rapidly decline towards deeper waters where bioerosion and carbonate accretion are slow and epibenthic/endolithic communities take years to mature. Accretion rates are highest in the lower euphotic zone (955 g/m2/yr), where the substrate is less prone to hydrodynamic force. Highest rates are found – inversely to bioerosion – on down-facing substrates, suggesting that bioerosion may be a key factor governing the preferential settlement and growth of calcareous epilithobionts on down-facing substrates. In context of a latitudinal gradient, the Azores carbonate cycling rates plot between known values from the cold-temperate Swedish Kosterfjord and the tropical Bahamas, with a total range of two orders in magnitude. Carbonate budget calculations for the bathymetrical transect yield a mean 266.9 kg of epilithic carbonate production, −54.6 kg of bioerosion, and 212.3 kg of annual net carbonate production per metre of coastline in the Azores C factory.

Description: Wide-angle seismic data have been used to determine the velocity and density structure of the crust and uppermost mantle beneath the Cape Verdes mid-plate swell. Seismic modelling reveals a ‘standard’ oceanic crust, ∼8 km in thickness, with no direct evidence for low-density bodies at the base of the crust. Gravity anomaly modelling within the constraints and resolution provided by the seismic model, does not preclude, however, a layer of crustal underplate up to 3 km thick beneath the swell crest. The modelling shows that while the seismically constrained crustal structure accounts for the short-wavelength free-air gravity anomaly, it fails to fully explain the long-wavelength anomaly. The main discrepancy is over the swell crest where the gravity anomaly, after correcting for crustal structure, is higher by about 30 mGal than it is over its flanks. The higher gravity can be explained if the top 100 km of the mantle beneath the swell crest is less dense than its surroundings by 30 kg m−3. The lack of evidence for low densities and velocities in the uppermost mantle, and high densities and velocities in the lower crust, suggests that neither a depleted swell root or crustal underplate are the origin of the observed shallower-than-predicted bathymetry and that, instead, the swell is most likely supported by dynamic uplift associated with an anomalously low density asthenospheric mantle.

Description: Aim: We investigate the relationship between local and regional richness in marine fouling assemblages using an expanded and globally replicated approach by incorporating two dimensions of diversity (taxonomic and functional) and different successional stages. Location: Global. Methods: In eight different biogeographic regions (Australia, Brazil, Chile, England, Italy, Japan, Portugal and Sweden) 68 polyvinylchloride (PVC) panels (15 × 15 × 0.3 cm) were deployed for colonization. Communities colonizing panels were analysed by measuring percentage cover at each of four different successional ages: 2, 4, 6 and 8 months. Local richness was assessed as the average number of species and functional groups (FGs) per panel and regional richness was evaluated as the estimated (Jack2) asymptote of the sample-accumulation curves for species and FG on experimental panels. Results: We found that the shape of the relationship between local and regional richness depended on successional stage and the type of richness considered, i.e. taxonomic or functional richness. Hardly any relationship was detectable between local taxonomic richness and regional taxonomic richness at any successional stage. In contrast, the relation between local functional and regional functional richness shows a unimodal pattern of change during succession, passing through the stages ‘independent’, ‘unsaturated rising’, ‘saturated rising’ and once again ‘independent’. Main conclusions: The relationship between local and regional richness, whether taxonomic or functional, frequently displays independence of the two scales, particularly in early and late phases of the successional process.

Description: A simple prognostic tool for gas hydrate (GH) quantification in marine sediments is presented based on a diagenetic transport-reaction model approach. One of the most crucial factors for the application of diagenetic models is the accurate formulation of microbial degradation rates of particulate organic carbon (POC) and the coupled formation of biogenic methane. Wallmann et al. (2006) suggested a kinetic formulation considering the ageing effects of POC and accumulation of reaction products (CH4, CO2) in the pore water. This model is applied to data sets of several ODP sites in order to test its general validity. Based on a thorough parameter analysis considering a wide range of environmental conditions, the POC accumulation rate (POCar in g/m2/yr) and the thickness of the gas hydrate stability zone (GHSZ in m) were identified as the most important and independent controls for biogenic GH formation. Hence, depth-integrated GH inventories in marine sediments (GHI in g of CH4 per cm2 seafloor area) can be estimated as: GHI=a ·POCar·GHSZb ·exp(−GHSZc/POCar/d)+e with a = 0.00214, b = 1.234, c = −3.339, d = 0.3148, e = −10.265. The transfer function gives a realistic first order approximation of the minimum GH inventory in low gas flux (LGF) systems. The overall advantage of the presented function is its simplicity compared to the application of complex numerical models, because only two easily accessible parameters need to be determined.

Description: Novel high-pressure biotechnical systems that were developed and applied for the study of anaerobic oxidation of methane (AOM) are described. The systems, referred to as high-pressure continuous incubation system (HP-CI system) and high-pressure manifold-incubation system (HP-MI system), allow for batch, fed-batch, and continuous gas-phase free incubation at high concentrations of dissolved methane and were designed to meet specific demands for studying environmental regulation and kinetics as well as for enriching microbial biomass in long-term incubation. Anoxic medium is saturated with methane in the first technical stage, and the saturated medium is supplied for biomass incubation in the second stage. Methane can be provided in continuous operation up to 20 MPa and the incubation systems can be operated during constant supply of gas-enriched medium at a hydrostatic pressure up to 45 MPa. To validate the suitability of the high-pressure systems, we present data from continuous and fed-batch incubation of highly active samples prepared from microbial mats from the Black Sea collected at a water depth of 213 m. In continuous operation in the HP-CI system initial methane-dependent sulfide production was enhanced 10- to 15-fold after increasing the methane partial pressure from near ambient pressure of 0.2 to 10.0 MPa at a hydrostatic pressure of 16.0 MPa in the incubation stage. With a hydraulic retention time of 14 h a stable effluent sulfide concentration was reached within less than 3 days and a continuing increase of the volumetric AOM rate from 1.2 to 1.7 mmol L−1 day−1 was observed over 14 days. In fed-batch incubation the AOM rate increased from 1.5 to 2.7 and 3.6 mmol L−1 day−1 when the concentration of aqueous methane was stepwise increased from 5 to 15 mmol L−1 and 45 mmol L−1. A methane partial pressure of 6 MPa and a hydrostatic pressure of 12 MPa in manifold fed-batch incubation in the HP-MI system yielded a sixfold increase in the volumetric AOM rate. Over subsequent incubation periods AOM rates increased from 0.6 to 1.2 mmol L−1 day−1 within 26 days of incubation. No inhibition of biomass activity was observed in all continuous and fed-batch incubation experiments. The organisms were able to tolerate high sulfide concentrations and extended starvation periods.

Description: Iron is a key micronutrient for phytoplankton growth in the surface ocean. Yet the significance of volcanism for the marine biogeochemical iron-cycle is poorly constrained. Recent studies, however, suggest that offshore deposition of airborne ash from volcanic eruptions is a way to inject significant amounts of bio-available iron into the surface ocean. Volcanic ash may be transported up to several tens of kilometers high into the atmosphere during large-scale eruptions and fine ash may stay aloft for days to weeks, thereby reaching even the remotest and most iron-starved oceanic regions. Scientific ocean drilling demonstrates that volcanic ash layers and dispersed ash particles are frequently found in marine sediments and that therefore volcanic ash deposition and iron-injection into the oceans took place throughout much of the Earth's history. Natural evidence and the data now available from geochemical and biological experiments and satellite techniques suggest that volcanic ash is a so far underestimated source for iron in the surface ocean, possibly of similar importance as aeolian dust. Here we summarise the development of and the knowledge in this fairly young research field. The paper covers a wide range of chemical and biological issues and we make recommendations for future directions in these areas. The review paper may thus be helpful to improve our understanding of the role of volcanic ash for the marine biogeochemical iron-cycle, marine primary productivity and the ocean-atmosphere exchange of CO2 and other gases relevant for climate in the Earth's history.

Description: Climate and weather extremes are sporadically recurring events that may have major local or regional impacts on the society and the environment. These events are typically related to unusually high or low temperature, prolonged dry or wet conditions, heavy precipitation, or extreme winds. Extreme events are part of the overall climate and weather alongside average conditions and variability, and thus are not unexpected as such. Climate change is expected to affect not only means but also variability and extremes. Some inferences can be based on past and present observations, but analyses of especially rare events are hampered by the availability of long time series. Over time, depending on how far the on-going global warming takes us from the present and the past climate conditions, the weather and climate statistics may well come to shift in ways that are well outside observational data. This may lead to shifts in frequency, intensity and geographical distribution of different extremes. Indeed, projected changes in some extremes over the 21st century are quite robust, such as generally increasing warm and decreasing cold extremes. Possible changes in some other aspects, for example storms, remain much more uncertain. Science-based information both on robust findings and on relevant uncertainties on changing extremes can provide useful information for sectorial planning, disaster risk prevention and overall reduction of societal vulnerability related to climate and weather

Description: In this overview, we consider the rationale and practice of invertebrate introductions into Lithuanian waters, the current distribution of introduced species, test the theoretical expectations of introduction of peracaridan species into lakes, and summarize the large-scale consequences of these translocations. After some early attempts to transfer locally available species within the country, peracaridan species from the Ponto-Caspian region were introduced into Lithuanian waters during the 1960s (Pontogammarus robustoides, Obesogammarus crassus, Chaetogammarus warpachowskyi, Paramysis lacustris, Limnomysis benedeni and Hemimysis anomala). After their establishment at the site of first introduction in a newly constructed water reservoir on the Nemunas River, further deliberate introductions into Lithuanian lakes and water reservoirs, as well as into more distant areas, were undertaken. These introduced species soon contributed to fish diet and a subsequent increase in the production of commercially important fish was envisaged. However, our collation of available data does not support the enhancement of fish production in Lithuanian lakes. Although perch (selected as a model fish species) assimilated the introduced species into diet and sometimes in large quantities, there was no subsequent influence on somatic growth rates when compared with perch from lakes devoid of alien peracaridans. Comparisons of littoral fish communities of different lakes, and commercial catches in the lake with the most numerous populations of introduced species also indicate no significant effect on fish production. Negative consequences of introduction are more than obvious in contrast. To date, most of the introduced Ponto-Caspian amphipod and mysid species occur in different sites of the Baltic Sea basin outside Lithuania, and their further dispersal may be expected. These species, especially amphipods, have proved to be highly ecologically aggressive, inducing change in resident macroinvertebrate communities and causing increase of biological contamination (i.e. detract from naturalness). Thus, the introduction of alien fodder species was ill advised from an economic, and even adverse from an environmental perspective.

Description: We review here the available information on methane (CH4) and nitrous oxide (N2O) from major marine, mostly coastal, oxygen (O2)-deficient zones formed both naturally and as a result of human activities (mainly eutrophication). Concentrations of both gases in subsurface waters are affected by ambient O2 levels to varying degrees. Organic matter supply to seafloor appears to be the primary factor controlling CH4 production in sediments and its supply to (and concentration in) overlying waters, with bottom-water O2-deficiency exerting only a modulating effect. High (micromolar level) CH4 accumulation occurs in anoxic (sulphidic) waters of silled basins, such as the Black Sea and Cariaco Basin, and over the highly productive Namibian shelf. In other regions experiencing various degrees of O2-deficiency (hypoxia to anoxia), CH4 concentrations vary from a few to hundreds of nanomolar levels. Since coastal O2-deficient zones are generally very productive and are sometimes located close to river mouths and submarine hydrocarbon seeps, it is difficult to differentiate any O2-deficiency-induced enhancement from in situ production of CH4 in the water column and its inputs through freshwater runoff or seepage from sediments. While the role of bottom-water O2-deficiency in CH4 formation appears to be secondary, even when CH4 accumulates in O2-deficient subsurface waters, methanotrophic activity severely restricts its diffusive efflux to the atmosphere. As a result, an intensification or expansion of coastal O2-deficient zones will probably not drastically change the present status where emission from the ocean as a whole forms an insignificant term in the atmospheric CH4 budget. The situation is different for N2O, the production of which is greatly enhanced in low-O2 waters, and although it is lost through denitrification in most suboxic and anoxic environments, the peripheries of such environments offer most suitable conditions for its production, with the exception of enclosed anoxic basins. Most O2-deficient systems serve as strong net sources of N2O to the atmosphere. This is especially true for coastal upwelling regions with shallow O2-deficient zones where a dramatic increase in N2O production often occurs in rapidly denitrifying waters. Nitrous oxide emissions from these zones are globally significant, and so their ongoing intensification and expansion is likely to lead to a significant increase in N2O emission from the ocean. However, a meaningful quantitative prediction of this increase is not possible at present because of continuing uncertainties concerning the formative pathways to N2O as well as insufficient data from key coastal regions.

Description: The role played by wind-forced anticyclones in the vertical transport and mixing at the ocean mesoscale is investigated with a primitive-equation numerical model in an idealized configuration. The focus of this work is to determine how the stratification impacts such transport. The flows, forced only at the surface by an idealized wind forcing, are predominantly horizontal and, on average, quasigeostrophic. Inside vortex cores and intense filaments, however, the dynamics is strongly ageostrophic. Mesoscale anticyclones appear as "islands" of increased penetration of wind energy into the ocean interior and they represent the maxima of available potential energy. The amount of available potential energy is directly correlated with the degree of stratification. The wind energy injected at the surface is transferred at depth through the generation and subsequent straining effect of Vortex Rossby Waves (VRWs), and through near-inertial internal oscillations trapped inside anticyclonic vortices. Both these mechanisms are affected by stratification. Stronger transfer but larger confinement close to the surface is found when the stratification is stronger. For weaker stratification, vertical mixing close to the surface is less intense but below about 150 m attains substantially higher values due to an increased contribution of both VRWs, whose time scale is on the order of few days, and of near-inertial motions, with a time scale of few hours.

Description: Whole genome amplification (WGA) approaches provide genomic information on single microbial cells and hold great promise for the field of environmental microbiology. Here, the microbial consortia of the marine sponge Aplysina aerophoba were sorted by fluorescence-activated cell sorting (FACS) and then subjected to WGA. A cosmid library was constructed from the WGA product of a sample containing two bacterial cells, one a member of the candidate phylum Poribacteria and one of a sponge-specific clade of Chloroflexi. Library screening led to the genomic characterization of three cosmid clones, encoding a polyketide synthase (PKS), a non-ribosomal peptide synthetase (NRPS) and the Chloroflexi 16S rRNA gene. PCR screening of WGA products from additional, FACS-sorted single bacterial symbiont cells supports the assignment of the Sup-PKS gene to the Poribacteria and the novel NRPS gene to the Chloroflexi. This promising single-cell genomics approach has permitted cloning of entire gene clusters from single microbial cells of known phylogenetic origin and thus provides a sought-after link between phylogeny and function.

Description: Das Phylum Schwämme und die mit ihnen assoziierten mikrobiellen Konsortien sind eine reichhaltige Quelle an neuartigen Wirkstoffen. Insbesondere die Naturstoffdiversität der Schwamm-assoziierten mikrobiellen Konsortien ist noch wenig untersucht. Methoden wie die klassische Kultivierung und die Metagenomik sind vielversprechende Ansätze zur umweltschonenden und nachhaltigen Gewinnung von neuen, bioaktiven Wirkstoffen aus Mikroorganismen. Insbesondere angesichts des vermehrten Auftretens von Antibiotika-resistenten Keimen sowie der ständigen Konfrontation mit neuen Infektionskrankheiten ist die Suche nach neuen Antibiotika/Antiinfektiva weiterhin dringend geboten.

Description: A brief description of the Deterministic Modelling Hydrological System (DMHS) ‘Runoff–Erosion–Pollution’ proposed by the first author is presented. This system is being developed with the aim of giving it a universal character so that it can be applied in mountainous and flat terrain, and in basins of different natural climatic zones regardless of their size. The main feature of the model is its independence of the scaling problem. The basis of our approach consists of a simple theory of runoff elements. This is different from the generally accepted use of partial differential equations such as the Saint Venant equation for surface and channel flow and the Boussinesq equation for underground waters describing the water movement from runoff formation origins to the basin outlet. The results of runoff simulation for six mountainous watersheds of different sizes across Eastern Siberia within the Lena River basin and their statistical evaluation are presented. The selected river basins ranged in size from about 40 km2 (small scale) to the entire Lena River basin (2·4 million km2), classified as a large-scale basin.

Description: Many marine sponges contain massive numbers of largely uncultivated, phylogenetically diverse bacteria that seem to be important contributors to the chemistry of these animals. Insights into the diversity, origin, distribution, and function of their metabolic gene communities are crucial to dissect the chemical ecology and biotechnological potential of sponge symbionts. This study reveals a sharp dichotomy between high and low microbial abundance sponges with respect to polyketide synthase (PKS) gene content, the presence of methyl-branched fatty acids, and the presence of members of the symbiotic candidate phylum "Poribacteria". For the symbiont-rich sponge Cacospongia mycofijiensis, a source of the tubulin-inhibiting fijianolides (=laulimalides), near-exhaustive large-scale sequencing of PKS gene-derived PCR amplicons was conducted. Although these amplicons exhibit high diversity at the sequence level, almost all of them belong to a single, architecturally unique group of PKSs present in "Poribacteria" and are proposed to synthesize methyl-branched fatty acids. Three components of this PKS were studied in vitro, providing initial insight into its biochemistry.

Description: Marine biologists have gone through a paradigm shift, from the assumption that marine populations are largely ‘open’ owing to extensive larval dispersal to the realization that marine dispersal is ‘more restricted than previously thought’. Yet, population genetic studies often reveal low levels of genetic structure across large geographic areas. On the other side, more direct approaches such as mark-recapture provide evidence of localized dispersal. To what extent can direct and indirect studies of marine dispersal be reconciled? One approach consists in applying genetic methods that have been validated with direct estimates of dispersal. Here, we use such an approach—genetic isolation by distance between individuals in continuous populations—to estimate the spatial scale of dispersal in five species of coral reef fish presenting low levels of genetic structure across the Caribbean. Individuals were sampled continuously along a 220-km transect following the Mesoamerican Barrier Reef, population densities were estimated from surveys covering 17 200 m2 of reef, and samples were genotyped at a total of 58 microsatellite loci. A small but positive isolation-by-distance slope was observed in the five species, providing mean parent-offspring dispersal estimates ranging between 7 and 42 km (CI 1–113 km) and suggesting that there might be a correlation between minimum/maximum pelagic larval duration and dispersal in coral reef fishes. Coalescent-based simulations indicate that these results are robust to a variety of dispersal distributions and sampling designs. We conclude that low levels of genetic structure across large geographic areas are not necessarily indicative of extensive dispersal at ecological timescales.

Description: Single crystal (U-Th)/He dating was applied to 24 apatite and 23 zircon grains from the Wetumpka impact structure, Alabama, USA. This small approximately 5–7.6 km impact crater was formed in a shallow marine environment, with no known preserved impact melt, thus offering a challenge to common geochronological techniques. A mean (U-Th)/He apatite and zircon age of 84.4 ± 1.4 Ma (2σ) was obtained, which is within error of the previously estimated Late Cretaceous impact age of approximately 83.5 Ma. In addition, helium diffusion modeling of apatite and zircon grains during fireball/contact, shock metamorphism, and hydrothermal events was undertaken, to show the influence of these individual thermal processes on resetting (U-Th)/He ages in the Wetumpka samples. This study has shown that the (U-Th)/He geochronological technique has real potential for dating impact structures, especially smaller and eroded impact structures that lack impact melt lithologies.

Description: The effects of different phases of semidiurnal and spring-neap tidal cycles on phytoplankton and environmental drivers were evaluated in a tidal, freshwater location of a mesotidal estuary (Guadiana estuary, SW Iberia). An Eulerian approach was used and sampling covered different seasons during 2008. Samples were collected during spring and neap tides, at high tide, mid-ebb, low tide and mid-flood. Several physical-chemical variables were measured, as well as phytoplankton abundance and biomass. Salinity was higher at high-tide and suspended particulate matter was higher during spring tides and flood, due to higher vertical mixing and resuspension of bottom sediments. Chlorophyll a concentration during winter and summer neap tides was higher than during spring tides, whilst the abundance of pennate diatoms was higher during winter and Spring spring tides than during neap tides, probably reflecting differences in river discharge. Overall, tidally-induced differences detected in the freshwater tidal reaches of the Guadiana estuary were not as considerable as those observed in the lower estuary. However, the occurrence of tidally-induced variability in some seasons reflects that thorough sampling programs to study estuarine tidal dynamics should be conducted throughout the year. Occasional sampling will not reflect the typical variability of these highly dynamic systems. (© 2010 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)

Description: The Baltic Sea is characterised by a heterogeneous oceanographic environment. The deep water layers forming the habitat of Baltic cod (Gadus morhua callarias L.) are subjected to frequently occurring pronounced anoxic conditions. Adverse oxygen conditions result in physiological stress for organisms living under these conditions. For cod e.g. a direct relationship between oxygen availability and food intake with a decreasing ingestion rate at hypoxia could be revealed. In the present study, the effects of oxygen deficiency on consumption rates were investigated and how these translate to stock size estimates in multi-species models. Based on results from laboratory experiments, a model was fitted to evacuation rates at different oxygen levels and integrated into the existing consumption model for Baltic cod. Individual mean oxygen corrected consumption rates were 0.1–10.9% lower than the uncorrected ones. At the currently low predator stock size, however, the effect of oxygen-reduced consumption on the total amount of eaten prey biomass and thus predation mortalities was only marginal. But should successful management lead to higher cod stock sizes in the future, then total predation mortalities will greatly increase and thus improved precision of these estimates would be valuable for the assessment of prey stocks.

Description: Parasitism is a common form of life and represents a strong selective pressure for host organisms. In response to this evolutionary pressure, vertebrates have developed genetically coded defences such as the major histocompatibility complex (MHC). Mechanisms of parasite-mediated selection not only maintain outstanding polymorphism in these genes but have also been proposed to further promote host population divergence and ultimately speciation because it can drive evolution of local adaptation in which MHC genes play a crucial role. This review first highlights the dynamics and complexity of parasite-mediated selection in natural systems, which not only depends on dominating parasite strategies and on the taxonomic diversity of the parasite community but also includes the differences in parasite communities between habitats and niches, creating divergent selection on locally adapted populations. Then the different ways in which MHC genes potentially allow vertebrates to respond to these dynamics and to adapt locally are outlined. Finally, it is proposed that varying selection strength in time and space may lead to variation in the strength of precopulatory reproductive isolation which has evolved to maintain local adaptation.

Description: We present the first application of Stochastic Heterogeneity Mapping based on the band-limited von Kármán function to a seismic reflection stack of a Mediterranean water eddy (meddy), a large salt lens of Mediterranean water. This process extracts two stochastic parameters directly from the reflectivity field of the seismic data: the Hurst number, which ranges from 0 to 1, and the correlation length (scale length). Lower Hurst numbers represent a richer range of high wavenumbers and correspond to a broader range of heterogeneity in reflection events. The Hurst number estimate for the top of the meddy (0.39) compares well with recent theoretical work, which required values between 0.25 and 0.5 to model internal wave surfaces in open ocean conditions based on simulating a Garrett-Munk spectrum (GM76) slope of −2. The scale lengths obtained do not fit as well to seismic reflection events as those used in other studies to model internal waves. We suggest two explanations for this discrepancy: (1) due to the fact that the stochastic parameters are derived from the reflectivity field rather than the impedance field the estimated scale lengths may be underestimated, as has been reported; and (2) because the meddy seismic image is a two-dimensional slice of a complex and dynamic three-dimensional object, the derived scale lengths are biased to the direction of flow. Nonetheless, varying stochastic parameters, which correspond to different spectral slopes in the Garrett-Munk spectrum (horizontal wavenumber spectrum), can provide an estimate of different internal wave scales from seismic data alone. We hence introduce Stochastic Heterogeneity Mapping as a novel tool in physical oceanography.

Description: We study the contribution of eastern-boundary density variations to sub-seasonal and seasonal anomalies of the strength and vertical structure of the Atlantic Meridional Overturning Circulation (AMOC) at 26.5° N, by means of the RAPID/MOCHA mooring array between April 2004 and October 2007. The major density anomalies are found in the upper 500 m, and they are often coherent down to 1400 m. The densities have 13-day fluctuations that are apparent down to 3500 m. The two strategies for measuring eastern-boundary density – a tall offshore mooring (EB1) and an array of moorings on the continental slope (EBH) – show little correspondence in terms of amplitude, vertical structure, and frequency distribution of the resulting basin-wide integrated transport fluctuations, implying that there are significant transport contributions between EB1 and EBH. Contrary to the original planning, measurements from EB1 cannot serve as backup or replacement for EBH: density needs to be measured directly at the continental slope to compute the full-basin density gradient. Fluctuations in density at EBH generate transport variability of 2 Sv rms in the AMOC, while the overall AMOC variability is 4.8 Sv rms. There is a pronounced deep-reaching seasonal cycle in density at the eastern boundary, which is apparent between 100 m and 1400 m, with maximum positive anomalies in spring and maximum negative anomalies in autumn. These changes drive anomalous southward upper mid-ocean flow in spring, implying maximum reduction of the AMOC, and vice-versa in autumn. The amplitude of the seasonal cycle of the AMOC arising from the eastern-boundary densities is 5.2 Sv peak-to-peak, dominating the 6.7 Sv peak-to-peak seasonal cycle of the total AMOC. Our analysis suggests that the seasonal cycle in density may be forced by the strong near-coastal seasonal cycle in wind stress curl.

Description: Summer heat waves have already resulted in mortality of coastal communities, including ecologically important seagrass meadows. Gene expression studies from controlled experiments can provide important insight as to how species/genotypes react to extreme events that will increase under global warming. In a common stress garden, we exposed three populations of eelgrass, Zostera marina, to extreme sea surface temperatures, simulating the 2003-European heat wave. Populations came from locations widely differing in their thermal regime, two northern European locations [Ebeltoft (Kattegat), Doverodde (Limfjord, Baltic Sea)], and one southern population from Gabicce Mare (Adriatic Sea), allowing to test for population specificity in the response to a realistic heat stress event. Eelgrass survival and growth as well as the expression of 12 stress associated candidate genes were assessed during and after the heat wave. Contrary to expectations, all populations suffered equally from 3 weeks of heat stress in terms of shoot loss. In contrast, populations markedly differed in multivariate measures of gene expression. While the gene expression profiles converged to pre-stress values directly after the heat wave, stress correlated genes were upregulated again 4 weeks later, in line with the observed delay in shoot loss. Target genes had to be selected based on functional knowledge in terrestrial plants, nevertheless, 10/12 genes were induced relative to the control treatment at least once during the heat wave in the fully marine plant Z. marina. This study underlines the importance of realistic stress and recovery scenarios in studying the impact of predicted climate change.

Description: We examined the influence of forecasted changes in global temperatures and pCO2 on N2 fixation and assimilation in the ecologically important cyanobacterium Trichodesmium spp. Changes of mRNA transcripts (nifH, glnA, hetR, psbA, psaB), protein (nitrogenase, glutamine synthetase) pools and enzymatic activity (nitrogenase) were measured under varying pCO2 and temperatures. High pCO2 shifted transcript patterns of all genes, resulting in a more synchronized diel expression. Under the same conditions, we did not observe any significant changes in the protein pools or in total cellular allocations of carbon and nitrogen (i.e. C : N ratio remained stable). Independently of temperature, high pCO2 (900 µatm) elevated N2 fixation rates. Levels of the key enzymes, nitrogenase and glutamine synthetase that mediate nitrogen assimilation did not increase, implying that the high pCO2 allowed higher reaction turnover rates through these key enzymes. Moreover, increased temperatures and high pCO2 resulted in higher C : P ratios. The plasticity in phosphorous stoichiometry combined with higher enzymatic efficiencies lead to higher growth rates. In cyanobacteria photosynthesis, carbon uptake, respiration, N2 fixation and nitrogen assimilation share cellular components. We propose that shifted cellular resource and energy allocation among those components will enable Trichodesmium grown at elevated temperatures and pCO2 to extend its niche in the future ocean, through both tolerance of a broader temperature range and higher P plasticity.

Description: Plankton sampling was conducted in the Baltic to obtain sprat larvae. Their individual drift patterns were back-calculated using a hydrodynamic model. The modelled positions along the individual drift trajectories were subsequently used to provide insight into the environmental conditions experienced by the larvae. Autocorrelation analysis revealed that successive otolith increment widths of individual larvae were not independent. Otolith increment width was then modelled using two different generalized additive model (GAM) analyses (with and without autocorrelation), using environmental variables determined for each modelled individual larval position as explanatory variables. The results indicate that otolith growth was not only influenced by the density of potential prey but was controlled by a number of simultaneously acting environmental factors. The final model, not considering autocorrelation, explained more than 80% of the variance of otolith growth, with larval age as a factor variable showing the strongest significant impact on otolith growth. Otolith growth was further explained by statistically significant ambient environmental factors such as temperature, bottom depth, prey density and turbulence. The GAM analysis, taking autocorrelation into account, explained almost 98% of the variability, with the previous otolith increment showing the strongest significant effect. Larval age as well as ambient temperature and prey abundance also had a significant effect. An alternative approach applied individual-based model (IBM) simulations on larval drift, feeding, growth and survival starting as exogenously feeding larvae at the backcalculated positions. The IBM results revealed optimal growth conditions for more than 97% of the larvae, with a tendency for our IBM to slightly overestimate larval growth.

Description: Water column data of carbon and carbon-relevant hydrographic and hydrochemical parameters from 188 cruises in the Arctic Mediterranean Seas, Atlantic and Southern Ocean have been retrieved and merged in a new data base: the CARINA (CARbon IN the Atlantic) Project. These data have gone through rigorous quality control (QC) procedures so as to improve the quality and consistency of the data as much as possible. Secondary quality control, which involved objective study of data in order to quantify systematic differences in the reported values, was performed for the pertinent parameters in the CARINA data base. Systematic biases in the data have been tentatively corrected in the data products. The products are three merged data files with measured, adjusted and interpolated data of all cruises for each of the three CARINA regions (Arctic Mediterranean Seas, Atlantic and Southern Ocean). Ninety-eight cruises were conducted in the "Atlantic" defined as the region south of the Greenland-Iceland-Scotland Ridge and north of about 30° S. Here we report the details of the secondary QC which was done on the total dissolved inorganic carbon (TCO2) data and the adjustments that were applied to yield the final data product in the Atlantic. Procedures of quality control – including crossover analysis between stations and inversion analysis of all crossover data – are briefly described. Adjustments were applied to TCO2 measurements for 17 of the cruises in the Atlantic Ocean region. With these adjustments, the CARINA data base is consistent both internally as well as with GLODAP data, an oceanographic data set based on the WOCE Hydrographic Program in the 1990s, and is now suitable for accurate assessments of, for example, regional oceanic carbon inventories, uptake rates and model validation.

Description: We examined the diel variation in nitrogen and carbon metabolism in Crocosphaera watsonii WH8501 at the physiological and gene expression level in order to determine the temporal constraints for N2 fixation and photosynthesis. N2 fixation and photosynthesis were restricted to the dark and light periods, respectively, during a 24 h light–dark cycle. All genes studied here except one (psbA2) showed diel variations in their expression levels. The highest variation was seen in nifH and nifX relative transcript abundance with a factor of 3–5 × 103 between light and dark periods. Photosynthesis genes showed less variation with a maximum factor of about 500 and always had high relative transcript abundances relative to other genes. At the protein level, the photosystems appeared more stable than the nitrogenase complex over a 24 h light–dark cycle, suggesting that C. watsonii retains the ability to photosynthesize during the dark period of the diel cycle. In contrast, nitrogenase is synthesized daily and exhibits peak abundance during the dark period. Our results have implications for field studies with respect to the interpretation of environmental gene expression data.

Description: The uptake of anthropogenic emission of carbon dioxide is resulting in a lowering of the carbonate saturation state and a drop in ocean pH. Understanding how marine calcifying organisms such as coralline algae may acclimatize to ocean acidification is important to understand their survival over the coming century. We present the first long-term perturbation experiment on the cold-water coralline algae, which are important marine calcifiers in the benthic ecosystems particularly at the higher latitudes. Lithothamnion glaciale, after three months incubation, continued to calcify even in undersaturated conditions with a significant trend towards lower growth rates with increasing pCO2. However, the major changes in the ultra-structure occur by 589 μatm (i.e. in saturated waters). Finite element models of the algae grown at these heightened levels show an increase in the total strain energy of nearly an order of magnitude and an uneven distribution of the stress inside the skeleton when subjected to similar loads as algae grown at ambient levels. This weakening of the structure is likely to reduce the ability of the alga to resist boring by predators and wave energy with severe consequences to the benthic community structure in the immediate future (50 years).

Description: The influence of various wind and wave conditions on the variability of downwelling irradiance Ed (490 nm) in water is subject of this study. The work is based on a two-dimensional Monte Carlo radiative transfer model with high spatial resolution. The model assumes conditions that are ideal for wave focusing, thus simulation results reveal the upper limit for light fluctuations. Local wind primarily determines the steepness of capillary-gravity waves which in turn dominate the irradiance variability near the surface. Down to 3 m depth, maximum irradiance peaks that exceed the mean irradiance Ed by a factor of more than 7 can be observed at low wind speeds up to 5 m s−1. The strength of irradiance fluctuations can be even amplified under the influence of higher ultra-gravity waves; thereby peaks can exceed 11 Ed. Sea states influence the light field much deeper; gravity waves can cause considerable irradiance variability even at 100 m depth. The simulation results show that under realistic conditions 50% radiative enhancements compared to the mean can still occur at 30 m depth. At greater depths, the underwater light variability depends on the wave steepness of the characteristic wave of a sea state; steeper waves cause stronger light fluctuations.

Description: The Arctic featured the strongest surface warming over the globe during the recent decades, and the temperature increase was accompanied by a rapid decline in sea ice extent. However, little is known about Arctic sea ice change during the Early Twentieth Century Warming (ETCW) during 1920–1940, also a period of a strong surface warming, both globally and in the Arctic. Here, we investigate the sensitivity of Arctic winter surface air temperature (SAT) to sea ice during 1875–2008 by means of simulations with an atmospheric general circulation model (AGCM) forced by estimates of the observed sea surface temperature (SST) and sea ice concentration. The Arctic warming trend since the 1960s is very well reproduced by the model. In contrast, ETCW in the Arctic is hardly captured. This is consistent with the fact that the sea ice extent in the forcing data does not strongly vary during ETCW. AGCM simulations with observed SST but fixed sea ice reveal a strong dependence of winter SAT on sea ice extent. In particular, the warming during the recent decades is strongly underestimated by the model, if the sea ice extent does not decline and varies only seasonally. This suggests that a significant reduction of Arctic sea ice extent may have also accompanied the Early Twentieth Century Warming, pointing toward an important link between anomalous sea ice extent and Arctic surface temperature variability.

Description: 2-D seismic data from the top and the western slope of Mergui Ridge in water depths between 300 and 2200 m off the Thai west coast have been investigated in order to identify mass transport deposits (MTDs) and evaluate the tsunamigenic potential of submarine landslides in this outer shelf area. Based on our newly collected data, 17 mass transport deposits have been identified. Minimum volumes of individual MTDs range between 0.3 km3 and 14 km3. Landslide deposits have been identified in three different settings: (i) stacked MTDs within disturbed and faulted basin sediments at the transition of the East Andaman Basin to the Mergui Ridge; (ii) MTDs within a pile of drift sediments at the basin-ridge transition; and (iii) MTDs near the edge of/on top of Mergui Ridge in relatively shallow water depths (〈 1000 m). Our data indicate that the Mergui Ridge slope area seems to have been generally unstable with repeated occurrence of slide events. We find that the most likely causes for slope instabilities may be the presence of unstable drift sediments, excess pore pressure, and active tectonics. Most MTDs are located in large water depths (〉 1000 m) and/or comprise small volumes suggesting a small tsunami potential. Moreover, the recurrence rates of failure events seem to be low. Some MTDs with tsunami potential, however, have been identified on top of Mergui Ridge. Mass-wasting events that may occur in the future at similar locations may trigger tsunamis if they comprise sufficient volumes. Landslide tsunamis, emerging from slope failures in the working area and affecting western Thailand coastal areas therefore cannot be excluded, though the probability is very small compared to the probability of earthquake-triggered tsunamis, arising from the Sunda Trench.

Description: Observations and model runs indicate trends in dissolved oxygen (DO) associated with current and ongoing global warming. However, a large-scale observation-to-model comparison has been missing and is presented here. This study presents a first global compilation of DO measurements covering the last 50 yr. It shows declining upper-ocean DO levels in many regions, especially the tropical oceans, whereas areas with increasing trends are found in the subtropics and in some subpolar regions. For the Atlantic Ocean south of 20° N, the DO history could even be extended back to about 70 yr, showing decreasing DO in the subtropical South Atlantic. The global mean DO trend between 50° S and 50° N at 300 dbar for the period 1960 to 2010 is –0.066 μmol kg−1 yr−1. Results of a numerical biogeochemical Earth system model reveal that the magnitude of the observed change is consistent with CO2-induced climate change. However, the pattern correlation between simulated and observed patterns of past DO change is negative, indicating that the model does not correctly reproduce the processes responsible for observed regional oxygen changes in the past 50 yr. A negative pattern correlation is also obtained for model configurations with particularly low and particularly high diapycnal mixing, for a configuration that assumes a CO2-induced enhancement of the C : N ratios of exported organic matter and irrespective of whether climatological or realistic winds from reanalysis products are used to force the model. Depending on the model configuration the 300 dbar DO trend between 50° S and 50° N is −0.027 to –0.047 μmol kg−1 yr−1 for climatological wind forcing, with a much larger range of –0.083 to +0.027 μmol kg−1 yr−1 for different initializations of sensitivity runs with reanalysis wind forcing. Although numerical models reproduce the overall sign and, to some extent, magnitude of observed ocean deoxygenation, this degree of realism does not necessarily apply to simulated regional patterns and the representation of processes involved in their generation. Further analysis of the processes that can explain the discrepancies between observed and modeled DO trends is required to better understand the climate sensitivity of oceanic oxygen fields and predict potential DO changes in the future.

Description: Ocean acidification and warming will be most pronounced in the Arctic Ocean. Aragonite shell-bearing pteropods in the Arctic are expected to be among the first species to suffer from ocean acidification. Carbonate undersaturation in the Arctic will first occur in winter and because this period is also characterized by low food availability, the overwintering stages of polar pteropods may develop into a bottleneck in their life cycle. The impacts of ocean acidification and warming on growth, shell degradation (dissolution), and mortality of two thecosome pteropods, the polar Limacina helicina and the boreal L. retroversa, were studied for the first time during the Arctic winter in the Kongsfjord (Svalbard). The abundance of L. helicina and L. retroversa varied from 23.5 to 120 ind m−2 and 12 to 38 ind m−2, and the mean shell size ranged from 920 to 981 μm and 810 to 823 μm, respectively. Seawater was aragonite-undersaturated at the overwintering depths of pteropods on two out of ten days of our observations. A 7-day experiment [temperature levels: 2 and 7 °C, pCO2 levels: 350, 650 (only for L. helicina) and 880 μatm] revealed a significant pCO2 effect on shell degradation in both species, and synergistic effects between temperature and pCO2 for L. helicina. A comparison of live and dead specimens kept under the same experimental conditions indicated that both species were capable of actively reducing the impacts of acidification on shell dissolution. A higher vulnerability to increasing pCO2 and temperature during the winter season is indicated compared with a similar study from fall 2009. Considering the species winter phenology and the seasonal changes in carbonate chemistry in Arctic waters, negative climate change effects on Arctic thecosomes are likely to show up first during winter, possibly well before ocean acidification effects become detectable during the summer season.

Description: The chemical characterization of filter high volume (HV) and Berner impactor (BI) samples PM during RHaMBLe (Reactive Halogens in the Marine Boundary Layer) 2007 shows that the Cape Verde aerosol particles are mainly composed of sea salt, mineral dust and associated water. Minor components are nss-salts, OC and EC. The influence from the African continent on the aerosol constitution was generally small but air masses which came from south-western Europe crossing the Canary Islands transported dust to the sampling site together with other loadings. The mean mass concentration was determined for PM10 to 17 μg/m3 from impactor samples and to 24.2 μg/m3 from HV filter samples. Non sea salt (nss) components of PM were found in the submicron fractions and nitrate in the coarse mode fraction. Bromide was found in all samples with much depleted concentrations in the range 1–8 ng/m3 compared to fresh sea salt aerosol indicating intense atmospheric halogen chemistry. Loss of bromide by ozone reaction during long sampling time is supposed and resulted totally in 82±12% in coarse mode impactor samples and in filter samples in 88±6% bromide deficits. A chloride deficit was determined to 8% and 1% for the coarse mode particles (3.5–10 μm; 1.2–3.5 μm) and to 21% for filter samples. During 14 May with high mineral dust loads also the maximum of OC (1.71μg/m3) and EC (1.25 μg/m3) was measured. The minimum of TC (0.25 μg/m3) was detected during the period 25 to 27 May when pure marine air masses arrived. The concentrations of carbonaceous material decrease with increasing particle size from 60% for the ultra fine particles to 2.5% in coarse mode PM. Total iron (dust vs. non-dust: 0.53 vs. 0.06 μg m3), calcium (0.22 vs. 0.03 μg m3) and potassium (0.33 vs. 0.02 μg m3) were found as good indicators for dust periods because of their heavily increased concentration in the 1.2 to 3.5 μm fraction as compared to their concentration during the non-dust periods. For the organic constituents, oxalate (78–151 ng/m3) and methanesulfonic acid (MSA, 25–100 ng/m3) are the major compounds identified. A good correlation between nss-sulphate and MSA was found for the majority of days indicating active DMS chemistry and low anthropogenic influences.

Description: A 2-Dimensional mathematical reaction-transport model was developed to study the impact of the mud-dwelling frenulate tubeworm Siboglinum sp. on the biogeochemistry of a sediment (MUC15) at the Captain Arutyunov mud volcano (CAMV). By explicitly describing the worm in its surrounding sediment, we are able to make budgets of processes occurring in- or outside of the worm, and to quantify how different worm densities and biomasses affect the anaerobic oxidation of methane (AOM) and sulfide reoxidation (HSox). The model shows that, at the observed densities, the presence of a thin worm body is sufficient to keep the upper 10 cm of sediment well homogenised with respect to dissolved substances, in agreement with observations. By this "bio-ventilation" activity, the worm pushes the sulfate-methane transition (SMT) zone downward to the posterior end of its body, and simultaneously physically separates the sulfide produced during the anaerobic oxidation of methane from oxygen. While there is little scope for AOM to take place in the tubeworm's body, 70% of the sulfide that is produced by sulfate reduction processes or that is advected in the sediment is preferentially shunted via the organism where it is oxidised by endosymbionts providing the energy for the worm's growth. The process of sulfide reoxidation, occurring predominantly in the worm's body is thus very distinct from the anaerobic oxidation of methane, which is a diffuse process that takes place in the sediments in the methane-sulfate transition zone. We show how the sulfide oxidation process is affected by increasing densities and length of the frenulates, and by upward advection velocity. Our biogeochemical model is one of the first to describe tubeworms explicitly. It can be used to directly link biological and biogeochemical observations at seep sites, and to study the impacts of mud-dwelling frenulates on the sediment biogeochemistry under varying environmental conditions. Also, it provides a tool to explore the competition between bacteria and fauna for available energy resources.

Description: We present a new tomographic model of the mantle in the area of the 2010 M8.8 Maule earthquake and surrounding regions. Increased ray coverage provided by the aftershock data allows us to image the detailed subducting slab structure in the mantle, from the region of flat slab subduction north of the Maule rupture to the area of overlapping rupture between the 1960 M9.5 and the 2010 M8.8 events to the south. We have combined teleseismic primary and depth phase arrivals with available local arrivals to better constrain the teleseismic earthquake locations in the region, which we use to conduct nested regional–global tomography. The new model reveals the detailed structure of the flat slab and its transition to a more moderately dipping slab in the Maule region. South of the Maule region, a steeply dipping relic slab is imaged from ∼200 to 1000 km depth that is distinct from the moderately dipping slab above it and from the more northerly slab at similar depths. We interpret the images as revealing both horizontal and vertical tearing of the slab at ∼38°S to explain the imaged pattern of slab anomalies in the southern portion of the model. In contrast, the transition from a horizontal to moderately subducting slab in the northern portion of the model is imaged as a continuous slab bend. We speculate that the tearing was most likely facilitated by a fracture zone in the downgoing plate or alternatively by a continental scale terrane boundary in the overriding plate.

Description: Global change drivers create new environmental scenarios and selective pressures, affecting plant species in various interacting ways. Plants respond with changes in phenology, physiology, and reproduction, with consequences for biotic interactions and community composition. We review information on phenotypic plasticity, a primary means by which plants cope with global change scenarios, recommending promising approaches for investigating the evolution of plasticity and describing constraints to its evolution. We discuss the important but largely ignored role of phenotypic plasticity in range shifts and review the extensive literature on invasive species as models of evolutionary change in novel environments. Plasticity can play a role both in the short-term response of plant populations to global change as well as in their long-term fate through the maintenance of genetic variation. In new environmental conditions, plasticity of certain functional traits may be beneficial (i.e., the plastic response is accompanied by a fitness advantage) and thus selected for. Plasticity can also be relevant in the establishment and persistence of plants in novel environments that are crucial for populations at the colonizing edge in range shifts induced by climate change. Experimental studies show taxonomically widespread plastic responses to global change drivers in many functional traits, though there is a lack of empirical support for many theoretical models on the evolution of phenotypic plasticity. Future studies should assess the adaptive value and evolutionary potential of plasticity under complex, realistic global change scenarios. Promising tools include resurrection protocols and artificial selection experiments.

Description: The future behaviour of the global ocean as a sink for CO2 is significant for climate change, but it is also important to understand its past by quantifying anthropogenic CO2 (Cant) in the ocean today. Unfortunately, this is complicated by the difficulty of deconvoluting Cant from the natural, unperturbed carbon cycle. Nonetheless, a range of techniques have been devised that perform this separation using the information implicit in other physical, biogeochemical and artificial ocean tracers. One such method is the TrOCA approach, whose parameterisation is derived from relationships between biogeochemical tracers within watermasses defined by age tracers such as CFC-11. TrOCA has a number of methodological advantages, and has been shown to be plausible, relative to other methods, in a number of studies. Here we examine the TrOCA approach by using it to deconvolute the known distribution of Cant from an ocean general circulation model (OGCM) simulation of the industrial period (1864–2004). TrOCA is evaluated at local, regional and global scales, with an emphasis on the wider applicability of the parameterisations derived at these scales. Our work finds that the published TrOCA parameterisation performs poorly when extrapolated beyond its calibration region, either with observational data or (especially) model output. Optimising TrOCA parameters using model output as a synthetic dataset leads to some small improvements, but the resulting TrOCA variants still perform poorly. Furthermore, there are large ranges on the optimised TrOCA parameters suggesting that a "universal" TrOCA parameterisation is not achieveable.

Description: Iron is a key limiting micro-nutrient for marine primary productivity. It can be supplied to the ocean by atmospheric dust deposition. Volcanic ash deposition into the ocean represents another external and so far largely neglected source of iron. This study demonstrates strong evidence for natural fertilisation in the iron-limited oceanic area of the NE Pacific, induced by volcanic ash from the eruption of Kasatochi volcano in August 2008. Atmospheric and oceanic conditions were favourable to generate a massive phytoplankton bloom in the NE Pacific Ocean which for the first time strongly suggests a connection between oceanic iron-fertilisation and volcanic ash supply.

Description: Very short-lived halocarbons are significant sources of reactive halogen in the marine boundary layer, and likely in the upper troposphere and lower stratosphere. Quantifying ambient concentrations in the surface ocean and atmosphere is essential for understanding the atmospheric impact of these trace gas fluxes. Despite the body of literature increasing substantially over recent years, calibration issues complicate the comparison of results and limit the utility of building larger-scale databases that would enable further development of the science (e.g. sea-air flux quantification, model validation, etc.). With this in mind, thirtyone scientists from both atmospheric and oceanic halocarbon communities in eight nations gathered in London in February 2008 to discuss the scientific issues and plan an international effort toward developing common calibration scales (http://tinyurl.com/c9cg58). Here, we discuss the outputs from this meeting, suggest the compounds that should be targeted initially, identify opportunities for beginning calibration and comparison efforts, and make recommendations for ways to improve the comparability of previous and future measurements.

Description: The main experimental approaches in aquatic community and ecosystem ecology will be presented along a gradient of scale: unenclosed field experiments, mesocosms with a natural mix of species, microcosms with an artificial mixture of species from cultures, and single species experiments in batch and continuous culture. Experimental manipulations usually consist of the addition or removal of (supposed) key organisms, addition of chemicals or alterations of the physical environment. Selecting the appropriate scale of experimentation is not only a question of technical and financial feasibility but also it has to consider the inevitable trade-offs between realism and control. Conclusions will be more widely accepted, if they are supported by experiments at a variety of scales. Key Concepts: The choice of the appropriate experimental scale has to face an inevitable trade-off between realism and control. Unenclosed field manipulations have the highest degree of realism but the least degree of control. Mesocosms operate with natural species assemblages, permit replication and control of experimental manipulations but are limited in temporal scale (weeks). Microcosm experiments operate with artificial communities assembled from cultures, offer a high degree of control, but lack the pre-history a mutual adaptation of the participating species. Experiments with single or few species rely on the 'model organism' concept. Small-scale experiments are biased against the detection of slow and space requiring processes.

Description: The conversion of fixed nitrogen to N2 in suboxic waters is estimated to contribute roughly a third to total oceanic losses of fixed nitrogen and is hence understood to be of major importance to global oceanic production and, therefore, to the role of the ocean as a sink of atmospheric CO2. At present heterotrophic denitrification and autotrophic anammox are considered the dominant sinks of fixed nitrogen. Recently, it has been suggested that the trophic nature of pelagic N2-production may have additional, "collateral" effects on the carbon cycle, where heterotrophic denitrification provides a shallow source of CO2 and autotrophic anammox a shallow sink. Here, we analyse the stoichiometries of nitrogen and associated carbon conversions in marine oxygen minimum zones (OMZ) focusing on heterotrophic denitrification, autotrophic anammox, and dissimilatory nitrate reduction to nitrite and ammonium in order to test this hypothesis quantitatively. For open ocean OMZs the combined effects of these processes turn out to be clearly heterotrophic, even with high shares of the autotrophic anammox reaction in total N2-production and including various combinations of dissimilatory processes which provide the substrates to anammox. In such systems, the degree of heterotrophy (ΔCO2:ΔN2), varying between 1.7 and 6, is a function of the efficiency of nitrogen conversion. On the contrary, in systems like the Black Sea, where suboxic N-conversions are supported by diffusive fluxes of NH4+ originating from neighbouring waters with sulphate reduction, much lower values of ΔCO2:ΔN2 can be found. However, accounting for concomitant diffusive fluxes of CO2, ratios approach higher values similar to those computed for open ocean OMZs. Based on our analysis, we question the significance of collateral effects concerning the trophic nature of suboxic N-conversions on the marine carbon cycle.

Description: Recent suggestions to slow down the increase in atmospheric carbon dioxide have included ocean fertilization by addition of the micronutrient iron to Southern Ocean surface waters, where a number of natural and artificial iron fertilization experiments have shown that low ambient iron concentrations limit phytoplankton growth. Using a coupled carbon-climate model with the marine biology's response to iron addition calibrated against data from natural iron fertilization experiments, we examine biogeochemical side effects of a hypothetical large-scale Southern Ocean Iron Fertilization (OIF) that need to be considered when attempting to account for possible OIF-induced carbon offsets. In agreement with earlier studies our model simulates an OIF-induced increase in local air-sea CO2 fluxes by about 60 GtC over a 100-year period, which amounts to about 40% of the OIF-induced increase in organic carbon export. Offsetting CO2 return fluxes outside the region and after stopping the fertilization at 1, 7, 10, 50, and 100 years are quantified for a typical accounting period of 100 years. For continuous Southern Ocean iron fertilization, the return flux outside the fertilized area cancels about 8% of the fertilization-induced CO2 air-sea flux within the fertilized area on a 100-yr timescale. This "leakage" effect has a similar radiative impact as the simulated enhancement of marine N2O emissions. Other side effects not yet discussed in terms of accounting schemes include a decrease in Southern Ocean oxygen levels and a simultaneous shrinking of tropical suboxic areas, and accelerated ocean acidification in the entire water column in the Southern Ocean on the expense of reduced globally averaged surface water acidification. A prudent approach to account for the OIF-induced carbon sequestration would account for global air-sea CO2 fluxes rather than for local fluxes into the fertilized area only. However, according to our model, this would underestimate the potential for offsetting CO2 emissions by about 20% on a 100 year accounting timescale. We suggest that a fair accounting scheme applicable to both terrestrial and marine carbon sequestration has to be based on emission offsets rather than on changes in individual carbon pools.

Description: Rising atmospheric CO2 is acidifying the surface ocean, a process which is expected to greatly influence the chemistry and biology of the future ocean. Following the development of iron-replete phytoplankton blooms in a coastal mesocosm experiment at 350, 700, and 1050 μatm pCO2, we observed significant increases in dissolved iron concentrations, Fe(II) concentrations, and Fe(II) half-life times during and after the peak of blooms in response to CO2 enrichment and concomitant lowering of pH, suggesting increased iron bioavailability. If applicable to the open ocean this may provide a negative feedback mechanism to the rising atmospheric CO2 by stimulating marine primary production.

Description: The response of the coccolithophore Emiliania huxleyi to rising CO2 concentrations is well documented for acclimated cultures where cells are exposed to the CO2 treatments for several generations prior to the experiment. The exact number of generations required for acclimation to CO2-induced changes in seawater carbonate chemistry, however, is unknown. Here we show that Emiliania huxleyi's short-term response (26 h) after cultures (grown at 500 μatm) were abruptly exposed to changed CO2 concentrations (~190, 410, 800 and 1500 μatm) is similar to that obtained with acclimated cultures under comparable conditions in earlier studies. Most importantly, from the lower CO2 levels (190 and 410 μatm) to 750 and 1500 μatm calcification decreased and organic carbon fixation increased within the first 8 to 14 h after exposing the cultures to changes in carbonate chemistry. This suggests that Emiliania huxleyi rapidly alters the rates of essential metabolical processes in response to changes in seawater carbonate chemistry, establishing a new physiological "state" (acclimation) within a matter of hours. If this relatively rapid response applies to other phytoplankton species, it may simplify interpretation of studies with natural communities (e.g. mesocosm studies and ship-board incubations), where often it is not feasible to allow for a pre-conditioning phase before starting experimental incubations.

Description: The physiological performance of two coccolithophore species, Emiliania huxleyi and Coccolithus braarudii, was investigated during long-term exposure to elevated pCO2 levels. Mono-specific cultures were grown over 152 (E. huxleyi) and 65 (C. braarudii) generations while pCO2 was gradually increased to maximum levels of 1150 μatm (E. huxleyi) and 930 μatm (C. braarudii) and kept constant thereafter. Rates of cell growth and cell quotas of particulate organic carbon (POC), particulate inorganic carbon (PIC) and total particulate nitrogen (TPN) were determined repeatedly throughout the incubation period. Increasing pCO2 caused a decrease in cell growth rate of 9% and 29% in E. huxleyi and C. braarudii, respectively. In both species cellular PIC:TPN and PIC:POC ratios decreased in response to rising pCO2, whereas no change was observed in the POC:TPN ratios of E. huxleyi and C. braarudii. These results are consistent with those obtained in shorter-term high CO2 exposure experiments following abrupt pertubations of the seawater carbonate system and indicate that for the strains tested here a gradual CO2 increase does not alleviate CO2/pH sensitivity.

Description: CO2/pH perturbation experiments were carried out under two different pCO2 levels (39.3 and 101.3 Pa) to evaluate effects of CO2-induced ocean acidification on the marine diatom Phaeodactylum tricornutum. After acclimation (〉20 generations) to ambient and elevated CO2 conditions (with corresponding pH values of 8.15 and 7.80, respectively), growth and photosynthetic carbon fixation rates of high CO2 grown cells were enhanced by 5% and 12%, respectively, and dark respiration stimulated by 34% compared to cells grown at ambient CO2. The half saturation constant (Km) for carbon fixation (dissolved inorganic carbon, DIC) increased by 20% under the low pH and high CO2 condition, reflecting a decreased affinity for HCO3– or/and CO2 and down-regulated carbon concentrating mechanism (CCM). In the high CO2 grown cells, the electron transport rate from photosystem II (PSII) was photoinhibited to a greater extent at high levels of photosynthetically active radiation, while non-photochemical quenching was reduced compared to low CO2 grown cells. This was probably due to the down-regulation of CCM, which serves as a sink for excessive energy. The balance between these positive and negative effects on diatom productivity will be a key factor in determining the net effect of rising atmospheric CO2 on ocean primary production.

Description: The potential impact of rising carbon dioxide (CO2) on carbon transfer from phytoplankton to bacteria was investigated during the 2005 PeECE III mesocosm study in Bergen, Norway. Sets of mesocosms, in which a phytoplankton bloom was induced by nutrient addition, were incubated under 1× (~350 μatm), 2× (~700 μatm), and 3× present day CO2 (~1050 μatm) initial seawater and sustained atmospheric CO2 levels for 3 weeks. 13C labelled bicarbonate was added to all mesocosms to follow the transfer of carbon from dissolved inorganic carbon (DIC) into phytoplankton and subsequently heterotrophic bacteria, and settling particles. Isotope ratios of polar-lipid-derived fatty acids (PLFA) were used to infer the biomass and production of phytoplankton and bacteria. Phytoplankton PLFA were enriched within one day after label addition, whilst it took another 3 days before bacteria showed substantial enrichment. Group-specific primary production measurements revealed that coccolithophores showed higher primary production than green algae and diatoms. Elevated CO2 had a significant positive effect on post-bloom biomass of green algae, diatoms, and bacteria. A simple model based on measured isotope ratios of phytoplankton and bacteria revealed that CO2 had no significant effect on the carbon transfer efficiency from phytoplankton to bacteria during the bloom. There was no indication of CO2 effects on enhanced settling based on isotope mixing models during the phytoplankton bloom, but this could not be determined in the post-bloom phase. Our results suggest that CO2 effects are most pronounced in the post-bloom phase, under nutrient limitation.

Description: The uptake of anthropogenic CO2 by the oceans has led to a rise in the oceanic partial pressure of CO2, and to a decrease in pH and carbonate ion concentration. This modification of the marine carbonate system is referred to as ocean acidification. Numerous papers report the effects of ocean acidification on marine organisms and communities but few have provided details concerning full carbonate chemistry and complementary observations. Additionally, carbonate system variables are often reported in different units, calculated using different sets of dissociation constants and on different pH scales. Hence the direct comparison of experimental results has been problematic and often misleading. The need was identified to (1) gather data on carbonate chemistry, biological and biogeochemical properties, and other ancillary data from published experimental data, (2) transform the information into common framework, and (3) make data freely available. The present paper is the outcome of an effort to integrate ocean carbonate chemistry data from the literature which has been supported by the European Network of Excellence for Ocean Ecosystems Analysis (EUR-OCEANS) and the European Project on Ocean Acidification (EPOCA). A total of 185 papers were identified, 100 contained enough information to readily compute carbonate chemistry variables, and 81 data sets were archived at PANGAEA – The Publishing Network for Geoscientific & Environmental Data. This data compilation is regularly updated as an ongoing mission of EPOCA.

Description: Ancient Lake Ohrid is a steep-sided, oligotrophic, karst lake that was tectonically formed most likely within the Pliocene and often referred to as a hotspot of endemic biodiversity. This study aims on tracing significant lake level fluctuations at Lake Ohrid using high-resolution acoustic data in combination with lithological, geochemical, and chronological information from two sediment cores recovered from sub-aquatic terrace levels at ca. 32 and 60m water depth. According to our data, significant lake level fluctuations with prominent lowstands of ca. 60 and 35m below the present water level occurred during Marine Isotope Stage (MIS) 6 and MIS 5, respectively. The effect of these lowstands on biodiversity in most coastal parts of the lake is negligible, due to only small changes in lake surface area, coastline, and habitat. In contrast, biodiversity in shallower areas was more severely affected due to disconnection of today sublacustrine springs from the main water body. Multichannel seismic data from deeper parts of the lake clearly image several clinoform structures stacked on top of each other. These stacked clinoforms indicate significantly lower lake levels prior to MIS 6 and a stepwise rise of water level with intermittent stillstands since its existence as water-filled body, which might have caused enhanced expansion of endemic species within Lake Ohrid.

Description: CO2 emissions are leading to an acidification of the oceans. Predicting marine community vulnerability towards acidification is difficult, as adaptation processes cannot be accounted for in most experimental studies. Naturally CO2 enriched sites thus can serve as valuable proxies for future changes in community structure. Here we describe a natural analogue site in the Western Baltic Sea. Seawater pCO2 in Kiel Fjord is elevated for large parts of the year due to upwelling of CO2 rich waters. Peak pCO2 values of 〉230 Pa (〉2300 μatm) and pHNBS values of 〈7.5 are encountered during summer and autumn, average pCO2 values are ~70 Pa (~700 μatm). In contrast to previously described naturally CO2 enriched sites that have suggested a progressive displacement of calcifying auto- and heterotrophic species, the macrobenthic community in Kiel Fjord is dominated by calcifying invertebrates. We show that blue mussels from Kiel Fjord can maintain control rates of somatic and shell growth at a pCO2 of 142 Pa (1400 μatm, pHNBS = 7.7). Juvenile mussel recruitment peaks during the summer months, when high water pCO2 values of ~100 Pa (~1000 μatm) prevail. Our findings indicate that calcifying keystone species may be able to cope with surface ocean pHNBS values projected for the end of this century when food supply is sufficient. However, owing to non-linear synergistic effects of future acidification and upwelling of corrosive water, peak seawater pCO2 in Kiel Fjord and many other productive estuarine habitats could increase to values 〉400 Pa (〉4000 μatm). These changes will most likely affect calcification and recruitment, and increase external shell dissolution.

Description: Gracilaria vermiculophylla (Ohmi) Papenf., an agar-producing red alga introduced from northeast Asia to Europe and North America, is often highly abundant in invaded areas. To assay its genetic diversity and identify the putative source of invasive populations, we analyzed the mitochondrial cytochrome c oxidase subunit I (cox1) gene from 312 individuals of G. vermiculophylla collected in 37 native and 32 introduced locations. A total of 19 haplotypes were detected: 17 in northeast Asia and three in Europe and eastern and western North America, with only one shared among all regions. The shared haplotype was present in all introduced populations and in ∼99% of individuals in the introduced areas. This haplotype was also found at three native locations in east Korea, west Japan, and eastern Russia. Both haplotype and nucleotide diversities were extremely low in Europe and North America compared to northeast Asia. Our study indicates that the East Sea/Sea of Japan is a likely donor region of the invasive populations of G. vermiculophylla in the east and west Atlantic and the east Pacific.

Description: Combined phylogenetic, physiological, and biochemical approaches revealed that differences in defense-related responses among 17 species belonging to the Gracilariaceae were consistent with their evolutionary history. An oxidative burst response resulting from activation of NADPH oxidase was always observed in two of the subgenera of Gracilaria sensu lato (Gracilaria, Hydropuntia), but not in Gracilariopsis and in species related to Gracilaria chilensis (“chilensis” clade). On the other hand, all species examined except Gracilaria tenuistipitata var. liui and Gracilariopsis longissima responded with up-regulation of agar oligosaccharide oxidase to an challenge with agar oligosaccharides. As indicated by pharmacological experiments conducted with Gracilaria chilensis and Gracilaria sp. “dura,” the up-regulation of agar oligosaccharide oxidase involved an NAD(P)H-dependent signaling pathway, but not kinase activity. By contrast, the activation of NADPH oxidase requires protein phosphorylation. Both responses are therefore independent, and the agar oligosaccharide-activated oxidative burst evolved after the capacity to oxidize agar oligosaccharide, probably providing additional defensive capacity to the most recently differentiated clades of Gracilariaceae. As demonstrated with Gracilaria gracilis, Gracilaria dura, and Gracilariopsis longissima, the different responses to agar oligosaccharides allow for a fast and nondestructive distinction among different clades of gracilarioids that are morphologically convergent. Based upon sequences of the chloroplast-encoded rbcL gene, this study suggests that at least some of the samples from NW America recorded as Gs. lemanaeiformis are probably Gs. chorda. Moreover, previous records of Gracilaria conferta from Israel are shown to be based upon misidentification of Gracilaria sp. “dura,” a species that belongs to the Hydropuntia subgenus.

Description: Ocean acidification, as a consequence of increasing marine pCO2, may have severe effects on the physiology of marine organisms. However, experimental studies remain scarce, in particular concerning fish. While adults will most likely remain relatively unaffected by changes in seawater pH, early life-history stages are potentially more sensitive – particularly the critical stage of fertilization, in which sperm motility plays a central role. In this study, the effects of ocean acidification (decrease of pHT to 7.55) on sperm motility of Baltic cod, Gadus morhua, were assessed. We found no significant effect of decreased pH on sperm speed, rate of change of direction or percent motility for the population of cod analyzed. We predict that future ocean acidification will probably not pose a problem for sperm behavior, and hence fertilization success, of Baltic cod.

Description: A shallow-water area in the Santa Barbara Channel, California, known collectively as the Coal Oil Point seep field, is one of the largest natural submarine hydrocarbon emission areas in the world. Both gas and oil are seeping constantly through a predominantly sandy seabed into the ocean. This study focused on the methanotrophic activity within the surface sediments (0–15 cm) of the permeable seabed in the so-called Brian Seep area at a water depth of ~10 m. Detailed investigations of the sediment biogeochemistry of active gas vents indicated that it is driven by fast advective transport of water through the sands, resulting in a deep penetration of oxidants (oxygen, sulfate). Maxima of microbial methane consumption were found at the sediment-water interface and in deeper layers of the sediment, representing either aerobic or anaerobic oxidation of methane, respectively. Methane consumption was relatively low (0.6–8.7 mmolm−2 d−1) in comparison to gas hydratebearing fine-grained sediments on the continental shelf. The low rates and the observation of free gas migrating through permeable coastal sediments indicate that a substantial proportion of methane can escape the microbial methane filter in coastal sediments.

Description: Two new brominated compounds, subereaphenol K (2) and 2-(3,5-dibromo-1-ethoxy-4-oxocyclohexa-2,5-dien-1-yl)acetamide (3), together with subereaphenol B (methyl 2-(2,4-dibromo-3,6-dihydroxyphenyl)acetate; 1) with a revised structure, and five dibromotyrosine-derived metabolites, 4–8, were isolated from the sponge Suberea sp. and characterized by 1D- and 2D-NMR spectroscopic and HR-MS spectrometric data. Compounds 1, 2, 6, and 8 exhibited various weak or moderate bioactivities, including antimicrobial and cytotoxic activities. Furthermore, compounds 1 and 2 inhibited human recombinant phosphodiesterase 4 (PDE4) with IC50 values of 2 μM, whereas compounds 6 and 8 were less active.

Description: Diet composition of the expanding southern species European anchovy Engraulis encrasicolus in the western Baltic Sea was investigated. Results revealed an interesting case of bentho-pelagic coupling with potential implications for local fish species through competition for food resources.

Description: Dimethylsulfoniopropionate (DMSP) is produced in surface seawater by phytoplankton. Phytoplankton culture experiments have shown that nanoeucaryotes (NANO) display much higher mean DMSP-to-Carbon or DMSP-to-Chlorophyll (Chl) ratios than Prochlorococcus (PRO), Synechococcus (SYN) or diatoms (DIAT). Moreover, the DMSP-lyase activity of algae which cleaves DMSP into dimethylsulfide (DMS) is even more group specific than DMSP itself. Ship-based observations have shown at limited spatial scales, that sea surface DMS-to-Chl ratios (DMS:Chl) are dependent on the composition of phytoplankton groups. Here we use satellite remote sensing of Chl (from SeaWiFS) and of Phytoplankton Group Dominance (PGD from PHYSAT) with ship-based sea surface DMS concentrations (8 cruises in total) to assess this dependence on an unprecedented spatial scale. PHYSAT provides PGD (either NANO, PRO, SYN, DIAT, Phaeocystis (PHAEO) or coccolithophores (COC)) in each satellite pixel (1/4° horizontal resolution). While there are identification errors in the PHYSAT method, it is important to note that these errors are lowest for NANO PGD which we typify by high DMSP:Chl. In summer, in the Indian sector of the Southern Ocean, we find that mean DMS:Chl associated with NANO + PHAEO and PRO + SYN + DIAT are 13.6±8.4 mmol g−1 (n=34) and 7.3±4.8 mmol g−1 (n=24), respectively. That is a statistically significant difference (P〈0.001) that is consistent with NANO and PHAEO being relatively high DMSP producers. However, in the western North Atlantic between 40° N and 60° N, we find no significant difference between the same PGD. This is most likely because coccolithophores account for the non-dominant part of the summer phytoplankton assemblages. Meridional distributions at 22° W in the Atlantic, and 95° W and 110° W in the Pacific, both show a marked drop in DMS:Chl near the equator, down to few mmol g−1, yet the basins exhibit different PGD (NANO in the Atlantic, PRO and SYN in the Pacific). In tropical and subtropical Atlantic and Pacific waters away from the equatorial and coastal upwelling, mean DMS:Chl associated with high and low DMSP producers are statistically significantly different, but the difference is opposite of that expected from culture experiments. Hence, in a majority of cases PGD is not of primary importance in controlling DMS:Chl variations. We therefore conclude that water-leaving radiance spectra obtained simultaneously from ocean color sensor measurements of Chl concentrations and dominant phytoplankton groups can not be used to predict global fields of DMS.

Description: Marine natural source electromagnetic data acquired on continental margins are often of considerable scientific and commercial interest. However, the large conductivity contrast between the ocean and coast causes this type of data to be severely distorted. For a 2-D coastal model, this distortion is most pronounced for the marine magnetotelluric and geomagnetic response function derived from induced currents flowing parallel to the coast. A maximal distortion occurs for a given period at a specific distance from the coast and causes severe anomalies in the magnitude and phase of the response functions. Based on a modelling study, we empirically relate the characteristic period and characteristic distance to physical parameters such as the ocean depth and the host resistivity. Based on a simple analytical approach, we test these approximations and show that maximum distortion occurs when destructive interference between the ocean and host response is at its highest. While the coast effect causes a large distortion in the marine responses we show through a resolution analysis that it does not mask subsurface conductivity anomalies but in fact increases the sensitivity to the seafloor.

Description: We present results of a seismic refraction experiment which determines the crustal and upper-mantle structure of an oceanic core complex (OCC) and its conjugate side located south of the 5°S ridge–transform intersection at the Mid-Atlantic Ridge. The core complex with a corrugated surface has been split by a change in location of active seafloor spreading, resulting in two massifs on either side of the current spreading axis. We applied a joint tomographic inversion of wide-angle reflected and refracted phases for five intersecting seismic profiles. The obtained velocity models are used to constrain the magmatic evolution of the core complex from the analysis of seismic layer 3 and crustal thickness. An abrupt increase of crustal velocities at shallow depth coincides with the onset of the seafloor corrugations at the exposed footwall. The observed velocity structure is consistent with the presence of gabbros directly beneath the corrugated fault surface. The thickness of the high-velocity body is constrained by PmP reflections to vary along and across axis between 〈3 and 5 km. The thickest crust is associated with the central phase of detachment faulting at the higher-elevated northern portion of the massif. Beneath the breakaway of the OCC the crust is 2.5 km thick and reveals significantly lower velocities. This implies that the fault initially exhumed low-velocity material overlying the gabbro plutons. In contrast, crust formed at the conjugate side during OCC formation is characterized by an up to 2-km-thick seismic layer 2 overlying a 1.7-km-thick seismic layer 3. Obtained upper-mantle velocities range from 7.3 to 7.9 km s−1 and seem to increase with distance from the median valley. However, velocities of 7.3–7.5 km s−1 beneath the older portions of the OCC may derive from deep fluid circulation and related hydrothermal alteration, which may likely be facilitated by the subsequent rifting. Our velocity models reveal a strongly asymmetric velocity structure across the ridge axis, associated with the accretion of gabbros into the footwall of the detachment fault and upper-crustal portions concentrated at the conjugate side. Our results do not support a substantial increase in the axial ridge's melt supply related to the final phase of detachment faulting. Hence, the footwall rifting at 5°S may be a generic mechanism of detachment termination under very low melt conditions, as predicted by recent numerical models of Tucholke et al.

Description: Anthropogenic emissions of nitrogen (N) to the atmosphere have been strongly increasing during the last century, leading to greater atmospheric N deposition to the oceans. The North Atlantic subtropical gyre (NASTG) is particularly impacted. Here, upwind sources of anthropogenic N from North American and European sources have raised atmospheric N deposition to rates comparable with N2 fixation in the gyre. However, the biogeochemical fate of the deposited N is unclear because there is no detectable accumulation in the surface waters. Most likely, deposited N accumulates in the main thermocline instead, where there is a globally unique pool of N in excess of the canonical Redfield ratio of 16 N:1 phosphorus (P). To investigate this depth zone as a sink for atmospheric N, we used a biogeochemical ocean transport model and year 2000 nutrient deposition data. We examined the maximum effects of three mechanisms that may transport excess N from the ocean surface to the main thermocline: physical transport, preferential P remineralization of sinking particles, and nutrient uptake and export by phytoplankton at higher than Redfield N:P ratios. Our results indicate that atmospheric deposition may contribute 13-19% of the annual excess N input to the main thermocline. Modeled nutrient distributions in the NASTG were comparable to observations only when non-Redfield dynamics were invoked. Preferential P remineralization could not produce realistic results on its own; if it is an important contributor to ocean biogeochemistry, it must co-occur with N2 fixation. The results suggest that: 1) the main thermocline is an important sink for anthropogenic N deposition, 2) non-Redfield surface dynamics determine the biogeochemical fate of atmospherically deposited nutrients, and 3) atmospheric N accumulation in the main thermocline has long term impacts on surface ocean biology.

Description: In the aftermath of an earthquake and tsunami on 11 March 2011 radioactive 137Cs was discharged from a damaged nuclear power plant to the sea off Fukushima Dai-ichi, Japan. Here we explore its dilution and fate with a state-of-the-art global ocean general circulation model, which is eddy-resolving in the region of interest. We find apparent consistency between our simulated circulation, estimates of 137Cs discharged ranging from 0.94 p Bq (Japanese Government, 2011) to 3.5 ± 0.7 p Bq (Tsumune et al., 2012), and measurements by Japanese authorities and the power plant operator. In contrast, our simulations are apparently inconsistent with the high 27 ± 15 p Bq discharge estimate of Bailly du Bois et al. (2012). Expressed in terms of a diffusivity we diagnose, from our simulations, an initial dilution on the shelf of 60 to 100 m2 s−1. The cross-shelf diffusivity is at 500 ± 300 m2 s−1 significantly higher and variable in time as indicated by its uncertainty. Expressed as an effective residence time of surface water on the shelf, the latter estimate transfers to 43 ± 16 days. As regards the fate of 137Cs, our simulations suggest that activities up to 4 mBq l−1 prevail in the Kuroshio-Oyashio Interfrontal Zone one year after the accident. This allows for low but detectable 0.1 to 0.3 m Bq l−1 entering the North Pacific Intermediate Water before the 137Cs signal is flushed away. The latter estimates concern the direct release to the sea only.

Description: This paper describes a 3-D lithospheric density model of the Western Continental Margin of India (WCMI) based on forward modelling of gravity data derived from satellite altimetry over the ocean and surface measurements on the Indian peninsula. The model covers the north-eastern Arabian Sea and the western part of the Indian Peninsula and incorporates constraints from a wide variety of geophysical and geological information. Salient features of the density model include: (1) the Moho depth varying from 13 km below the oceanic crust to 46 km below the continental interior; (2) the lithosphere–asthenosphere boundary (LAB) located at depths between 70 km in the southwestern corner (under oceanic crust) and about 165 km below the continental region; (3) thickening of the crust under the Chagos–Laccadive and Laxmi Ridges and (4) a revised definition of the continent–ocean boundary. The 3-D density structure of the region enables us to propose an evolutionary model of the WCMI that revisits earlier views of passive rifting. The first stage of continental-scale rifting of Madagascar from India at about 90 Ma is marked by relatively small amounts of magmatism. A second episode of rifting and large-scale magmatism was possibly initiated around 70 Ma with the opening of the Gop Rift. Subsequently at around 68 Ma, the drifting away of the Seychelles and formation of the Laxmi Ridge was a consequence of the down-faulting of the northern margin. During this second episode of rifting, the northern part of the WCMI witnessed massive volcanism attributed to interaction with the Reunion hotspot at around 65 Ma. Subsequent stretching of the transitional crust between about 65 and 62 Ma formed the Laxmi Basin, the southward extension of the failed Gop Rift. As the interaction between plume and lithosphere continued, the Chagos–Laccadive Ridge was emplaced on the edge of the nascent oceanic crust/rifted continental margin in the south as the Indian Plate was moving northwards.

Description: The global Late Pliocene/Early Pleistocene cooling (~3.0–2.0 million years ago – Ma) concurred with extremely high diatom and biogenic opal production in most of the major coastal upwelling regions. This phenomenon was particularly pronounced in the Benguela upwelling system (BUS), off Namibia, where it is known as the Matuyama Diatom Maximum (MDM). Our study focuses on a new diatom silicon isotope (δ30Si) record covering the MDM in the BUS. Unexpectedly, the variations in δ30Si signal follow biogenic opal content, whereby the highest δ30Si values correspond to the highest biogenic opal content. We interpret the higher δ30Si values during the MDM as a result of a stronger degree of silicate utilisation in the surface waters caused by high productivity of mat-forming diatom species. This was most likely promoted by weak upwelling intensity dominating the BUS during the Late Pliocene/Early Pleistocene cooling combined with a large silicate supply derived from a strong Southern Ocean nutrient leakage responding to the expansion of Antarctic ice cover and the resulting stratification of the polar ocean 3.0–2.7 Ma ago. A similar scenario is hypothesized for other major coastal upwelling systems (e.g. off California) during this time interval, suggesting that the efficiency of the biological carbon pump was probably sufficiently enhanced in these regions during the MDM to have significantly increased the transport of atmospheric CO2 to the deep ocean. In addition, the coeval extension of the area of surface water stratification in both the Southern Ocean and the North Pacific, which decreased CO2 release to the atmosphere, led to further enhanced atmospheric CO2 drawn-down and thus contributed significantly to Late Pliocene/Early Pleistocene cooling.

Description: The cuttlefish Sepia officinalis expresses several hemocyanin isoforms with potentially different pH optima, indicating their reliance on efficient pH regulation in the blood. Ongoing ocean warming and acidification could influence the oxygen-binding properties of respiratory pigments in ectothermic marine invertebrates. This study examined whether S. officinalis differentially expresses individual hemocyanin isoforms to maintain optimal oxygen transport during development and acclimation to elevated seawater pCO2 and temperature. Using quantitative PCR, we measured relative mRNA expression levels of three different hemocyanin isoforms in several ontogenetic stages (embryos, hatchlings, juveniles, and adults), under different temperatures and elevated seawater pCO2. Our results indicate moderately altered hemocyanin expression in all embryonic stages acclimated to higher pCO2, while hemocyanin expression in hatchlings and juveniles remained unaffected. During the course of development, total hemocyanin expression increased independently of pCO2 or thermal acclimation status. Expression of isoform 3 is reported for the first time in a cephalopod in this study and was found to be generally low but highest in the embryonic stages (0.2% of total expression). Despite variable hemocyanin expression, hemolymph total protein concentrations remained constant in the experimental groups. Our data provide first evidence that ontogeny has a stronger influence on hemocyanin isoform expression than the environmental conditions chosen, and they suggest that hemocyanin protein abundance in response to thermal acclimation is regulated by post-transcriptional/translational rather than by transcriptional modifications

Description: It is essential to predict the impact of elevated Pco2 on marine organisms and habitats to anticipate the severity and consequences of future ocean chemistry change. Despite the importance of carry-over effects in the evolutionary history of marine organisms, few studies have considered links between life-history stages when determining how marine organisms will respond to elevated Pco2, and none have considered the link between adults and their offspring. Herein, we exposed adults of wild and selectively bred Sydney rock oysters, Saccostrea glomerata to elevated Pco2 during reproductive conditioning and measured the development, growth and survival response of their larvae. We found that elevated Pco2 had a negative impact on larvae of S. glomerata causing a reduction in growth, rate of development and survival. Exposing adults to elevated Pco2 during reproductive conditioning, however, had positive carry-over effects on larvae. Larvae spawned from adults exposed to elevated Pco2 were larger and developed faster, but displayed similar survival compared with larvae spawned from adults exposed to ambient Pco2. Furthermore, selectively bred larvae of S. glomerata were more resilient to elevated Pco2 than wild larvae. Measurement of the standard metabolic rate (SMR) of adult S. glomerata showed that at ambient Pco2, SMR is increased in selectively bred compared with wild oysters and is further increased during exposure to elevated Pco2. This study suggests that sensitive marine organisms may have the capacity to acclimate or adapt to elevated Pco2 over the next century and a change in energy turnover indicated by SMR may be a key process involved.

Description: Although crucial for the understanding of adaptive evolution, genetically resolved examples of local adaptation are rare. To maximize survival and reproduction in their local environment, hosts should resist their local parasites and pathogens. The major histocompatibility complex (MHC) with its key function in parasite resistance represents an ideal candidate to investigate parasite-mediated local adaptation. Using replicated field mesocosms, stocked with second-generation lab-bred three-spined stickleback hybrids of a lake and a river population, we show local adaptation of MHC genotypes to population-specific parasites, independently of the genetic background. Increased allele divergence of lake MHC genotypes allows lake fish to fight the broad range of lake parasites, whereas more specific river genotypes confer selective advantages against the less diverse river parasites. Hybrids with local MHC genotype gained more body weight and thus higher fitness than those with foreign MHC in either habitat, suggesting the evolutionary significance of locally adapted MHC genotypes.

Description: Oxygen-deficient waters in the ocean, generally referred to as oxygen minimum zones (OMZ), are expected to expand as a consequence of global climate change. Poor oxygenation is promoting microbial loss of inorganic nitrogen (N) and increasing release of sediment-bound phosphate (P) into the water column. These intermediate water masses, nutrient-loaded but with an N deficit relative to the canonical N:P Redfield ratio of 16:1, are transported via coastal upwelling into the euphotic zone. To test the impact of nutrient supply and nutrient stoichiometry on production, partitioning and elemental composition of dissolved (DOC, DON, DOP) and particulate (POC, PON, POP) organic matter, three nutrient enrichment experiments were conducted with natural microbial communities in shipboard mesocosms, during research cruises in the tropical waters of the southeast Pacific and the northeast Atlantic. Maximum accumulation of POC and PON was observed under high N supply conditions, indicating that primary production was controlled by N availability. The stoichiometry of microbial biomass was unaffected by nutrient N:P supply during exponential growth under nutrient saturation, while it was highly variable under conditions of nutrient limitation and closely correlated to the N:P supply ratio, although PON:POP of accumulated biomass generally exceeded the supply ratio. Microbial N:P composition was constrained by a general lower limit of 5:1. Channelling of assimilated P into DOP appears to be the mechanism responsible for the consistent offset of cellular stoichiometry relative to inorganic nutrient supply and nutrient drawdown, as DOP build-up was observed to intensify under decreasing N:P supply. Low nutrient N:P conditions in coastal upwelling areas overlying O2-deficient waters seem to represent a net source for DOP, which may stimulate growth of diazotrophic phytoplankton. These results demonstrate that microbial nutrient assimilation and partitioning of organic matter between the particulate and the dissolved phase are controlled by the N:P ratio of upwelled nutrients, implying substantial consequences for nutrient cycling and organic matter pools in the course of decreasing nutrient N:P stoichiometry.

Description: Oceanic emissions of halogenated very short-lived substances (VSLS) are expected to contribute significantly to the stratospheric halogen loading and therefore to ozone depletion. The amount of VSLS transported into the stratosphere is estimated based on in-situ observations around the tropical tropopause layer (TTL) and on modeling studies which mostly use prescribed global emission scenarios to reproduce observed atmospheric concentrations. In addition to upper-air VSLS measurements, direct observations of oceanic VSLS emissions are available along ship cruise tracks. Here we use such in-situ observations of VSLS emissions from the West Pacific and tropical Atlantic together with an atmospheric Lagrangian transport model to estimate the direct contribution of bromoform (CHBr3), and dibromomethane (CH2Br2) to the stratospheric bromine loading as well as their ozone depletion potential. Our emission-based estimates of VSLS profiles are compared to upper-air observations and thus link observed oceanic emissions and in situ TTL measurements. This comparison determines how VSLS emissions and transport in the cruise track regions contribute to global upper-air VSLS estimates. The West Pacific emission-based profiles and the global upper-air observations of CHBr3 show a relatively good agreement indicating that emissions from the West Pacific provide an average contribution to the global CHBr3 budget. The tropical Atlantic, although also being a CHBr3 source region, is of less importance for global upper-air CHBr3 estimates as revealed by the small emission-based abundances in the TTL. Western Pacific CH2Br2 emission-based estimates are considerably smaller than upper-air observations as a result of the relatively low sea-to-air flux found in the West Pacific. Together, CHBr3 and CH2Br2 emissions from the West Pacific are projected to contribute to the stratospheric bromine budget with 0.4 pptv Br on average and 2.3 pptv Br for cases of maximum emissions through product and source gas injection. These relatively low estimates reveal that the tropical West Pacific, although characterized by strong convective transport, might overall contribute less VSLS to the stratospheric bromine budget than other regions as a result of only low CH2Br2 and moderate CHBr3 oceanic emissions.

Description: In this study we present an initial dataset of Mn/Ca and Fe/Ca ratios in tests of benthic foraminifera from the Peruvian oxygen minimum zone (OMZ) determined with SIMS. These results are a contribution to a better understanding of the proxy potential of these elemental ratios for ambient redox conditions. Foraminiferal tests are often contaminated by diagenetic coatings, like Mn rich carbonate- or Fe and Mn rich (oxyhydr)oxide coatings. Thus, it is substantial to assure that the cleaning protocols are efficient or that spots chosen for microanalyses are free of contaminants. Prior to the determination of the element/Ca ratios, the distributions of several elements (Ca, Mn, Fe, Mg, Ba, Al, Si, P and S) in tests of the shallow infaunal species Uvigerina peregrina and Bolivina spissa were mapped with an electron microprobe (EMP). To visualize the effects of cleaning protocols uncleaned and cleaned specimens were compared. The cleaning protocol included an oxidative cleaning step. An Fe rich phase was found on the inner test surface of uncleaned U. peregrina specimens. This phase was also enriched in Al, Si, P and S. A similar Fe rich phase was found at the inner test surface of B. spissa. Specimens of both species treated with oxidative cleaning show the absence of this phase. Neither in B. spissa nor in U. peregrina were any hints found for diagenetic (oxyhydr)oxide or carbonate coatings. Mn/Ca and Fe/Ca ratios of single specimens of B. spissa from different locations have been determined by secondary ion mass spectrometry (SIMS). Bulk analyses using solution ICP-MS of several samples were compared to the SIMS data. The difference between SIMS analyses and ICP-MS bulk analyses from the same sampling sites was 14.0–134.8 μmol mol−1 for the Fe/Ca and 1.68(±0.41) μmol mol−1 for the Mn/Ca ratios. This is in the same order of magnitude as the variability inside single specimens determined with SIMS at these sampling sites (1σ[Mn/Ca] = 0.35–2.07 μmol mol−1; 1σ[Fe/Ca] = 93.9–188.4 μmol mol−1). The Mn/Ca ratios in the calcite were generally relatively low (2.21–9.93 μmol mol−1) but in the same magnitude and proportional to the surrounding pore waters (1.37–6.67 μmol mol−1). However, the Fe/Ca ratios in B. spissa show a negative correlation to the concentrations in the surrounding pore waters. Lowest foraminiferal Fe/Ca ratios (87.0–101.0 μmol mol−1) were found at 465 m water depth, a location with a strong sharp Fe peak in the pore water next to the sediment surface and respectively, high Fe concentrations in the surrounding pore waters. Previous studies found no living specimens of B. spissa at this location. All these facts hint that the analysed specimens already were dead before the Fe flux started and the sampling site just recently turned anoxic due to fluctuations of the lower boundary of the OMZ near the sampling site (465 m water depth). Summarized Mn/Ca and Fe/Ca ratios are potential proxies for redox conditions, if cleaning protocols are carefully applied. The data presented here may be rated as base for the still pending detailed calibration.

Description: In industrialized and/or agriculturally used landscapes, inhabiting species are exposed to a variety of anthropogenic changes in their environments. Genetic diversity may be reduced if populations encounter founder events, bottlenecks, or isolation. Conversely, genetic diversity may increase if populations adapt to changes in selective regimes in newly created habitats. With the present study, genetic variability of 918 sticklebacks from 43 samplings (21.3 ± 3.8 per sample) at 36 locations from cultivated landscapes in Northwest Germany was analyzed at nine neutral microsatellite loci. To test if differentiation is influenced by habitat alterations, sticklebacks were collected from ancient running waters and adjacent artificial stagnant waters, from brooks with salt water inflow of anthropogenic and natural origin and adjacent freshwater sites. Overall population structure was dominated by isolation by distance (IBD), which was significant across all populations, and analysis of molecular variance (AMOVA) revealed that 10.6% of the variation was explained by river catchment area. Populations in anthropogenic modified habitats deviated from the general IBD structure and in the AMOVA, grouping by habitat type running/stagnant water explained 4.9% of variation and 1.4% of the variation was explained by salt-/freshwater habitat. Sticklebacks in salt-polluted water systems seem to exhibit elevated migratory activity between fresh- and saltwater habitats, reducing IBD. In other situations, populations showed distinct signs of genetic isolation, which in some locations was attributed to mechanical migration barriers, but in others to potential anthropogenic induced bottleneck or founder effects. The present study shows that anthropogenic habitat alterations may have diverse effects on the population genetic structure of inhabiting species. Depending on the type of habitat change, increased genetic differentiation, diversification, or isolation are possible consequences.

Description: The aim of this study is to determine cloud-type resolved cloud radiative budgets and cloud radiative effects from surface measurements of broadband radiative fluxes over the Atlantic Ocean. Furthermore, based on simultaneous observations of the state of the cloudy atmosphere, a radiative closure study has been performed by means of the ECHAM5 single column model in order to identify the model's ability to realistically reproduce the effects of clouds on the climate system. An extensive database of radiative and atmospheric measurements has been established along five meridional cruises of the German research icebreaker Polarstern. Besides pyranometer and pyrgeometer for downward broadband solar and thermal radiative fluxes, a sky imager and a microwave radiometer have been utilized to determine cloud fraction and cloud type on the one hand and temperature and humidity profiles as well as liquid water path for warm non-precipitating clouds on the other hand. Averaged over all cruise tracks, we obtain a total net (solar + thermal) radiative flux of 144 W m(-2) that is dominated by the solar component. In general, the solar contribution is large for cirrus clouds and small for stratus clouds. No significant meridional dependencies were found for the surface radiation budgets and cloud effects. The strongest surface longwave cloud effects were shown in the presence of low level clouds. Clouds with a high optical density induce strong negative solar radiative effects under high solar altitudes. The mean surface net cloud radiative effect is -33 W m(-2). For the purpose of quickly estimating the mean surface longwave, shortwave and net cloud effects in moderate, subtropical and tropical climate regimes, a new parameterisation was created, considering the total cloud amount and the solar zenith angle. The ECHAM5 single column model provides a surface net cloud effect that is more cooling by 17 W m(-2) compared to the radiation observations. This overestimation in solar cooling is mostly caused by the shortwave impact of convective clouds. The latter show a large overestimation in solar cooling of up to 114 W m(-2). Mean cloud radiative effects of cirrus and stratus clouds were simulated close to the observations.

Description: The sea-surface microlayer (SML) is located within the boundary between the atmosphere and hydrosphere. The high spatial and temporal variability of the SML's properties, however, have hindered a clear understanding of interactions between biotic and abiotic parameters at or across the air-water interface. Among the factors changing the physical and chemical environment of the SML, wind speed is an important one. In order to examine the temporal effects of minimized wind influence, SML samples were obtained from the coastal zone of the southern Baltic Sea and from mesocosm experiments in a marina to study naturally and artificially calmed sea surfaces. Organic matter concentrations as well as abundance, (3)H-thymidine incorporation, and the community composition of bacteria in the SML (bacterioneuston) compared to the underlying bulk water (ULW) were analyzed. In all SML samples, dissolved organic carbon and nitrogen were only slightly enriched and showed low temporal variability, whereas particulate organic carbon and nitrogen were generally greatly enriched and highly variable. This was especially pronounced in a dense surface film (slick) that developed during calm weather conditions as well as in the artificially calmed mesocosms. Overall, bacterioneuston abundance and productivity correlated with changing concentrations of particulate organic matter. Moreover, changes in the community composition in the field study were stronger in the particle-attached than in the non-attached bacterioneuston. This implies that decreasing wind enhances the importance of particle-attached assemblages and finally induces a succession of the bacterial community in the SML. Eventually, under very calm meteorological conditions, there is an uncoupling of the bacterioneuston from the ULW.

Description: The internal consistency of measurements and computations of components of the CO2-system, namely total alkalinity (AT), total dissolved carbon dioxide (CT), CO2 fugacity (fCO2), and pH, has been confirmed repeatedly in open ocean studies when the CO2 system had been over determined. Differences between measured and computed properties, such as ΔfCO2 (=fCO2(measured) – fCO2(computed from AT and CT))/ fCO2(measured)× 100), there are usually below 5%. Recently, Hoppe et al. (2010) provided evidence of significantly larger ΔfCO2 in experimental setups. These observations are currently not well understood. Here we discuss a case from a series of phytoplankton culture experiments with ΔfCO2 of up to about 25%. ΔfCO2 varied systematically during the course of these experiments and showed a clear correlation with the accumulation of dissolved organic carbon (DOC). Culture and mesocosm experiments are often carried out under very high initial nutrient concentrations, yielding high biomass concentrations that in turn often lead to a substantial build-up of DOC. DOC can reach concentrations much higher than typically observed in the open ocean. To the extent that DOC includes organic acids and bases, it will contribute to the alkalinity of the seawater contained in the experimental device. Our analysis suggests that whenever substantial amounts of DOC are produced during the experiment, standard computer programs used to compute CO2 fugacity can underestimate true fCO2 significantly when the computation is based on AT and CT. Alternative explanations for large ΔfCO2, e.g. uncertainties of pKs, are explored as well, but are found to be of minor importance. Unless the effect of DOC-alkalinity is accounted for, this might lead to significant errors in the interpretation of the system under consideration to the experimentally applied CO2 perturbation, which could misguide the development of parameterisations used in simulations with global carbon cycle models in future CO2-scenarios.

Description: Emiliania huxleyi (strain B 92/11) was exposed to different nutrient supply, CO2 and temperature conditions in phosphorus controlled chemostats to investigate effects on organic carbon exudation and partitioning between the pools of particulate organic carbon (POC) and dissolved organic carbon (DOC). 14C incubation measurements for primary production (PP) and extracellular release (ER) were performed. Chemical analysis included the amount and composition of high molecular weight (〉1 kDa) dissolved combined carbohydrates (HMW-dCCHO), particulate combined carbohydrates (pCCHO) and the carbon content of transparent exopolymer particles (TEP-C). Applied CO2 and temperature conditions were 300, 550 and 900 μatm pCO2 at 14 °C, and additionally 900 μatm pCO2 at 18 °C simulating a greenhouse ocean scenario. Enhanced nutrient stress by reducing the dilution rate (D) from D = 0.3 d−1 to D = 0.1 d−1 (D = μ) induced the strongest response in E. huxleyi. At μ = 0.3 d−1, PP was significantly higher at elevated CO2 and temperature and DO14C production correlated to PO14C production in all treatments, resulting in similar percentages of extracellular release (PER; (DO14C production/PP) × 100) averaging 3.74 ± 0.94%. At μ = 0.1 d−1, PO14C production decreased significantly, while exudation of DO14C increased. Thus, indicating a stronger partitioning from the particulate to the dissolved pool. Maximum PER of 16.3 ± 2.3% were observed at μ = 0.1 d−1 at elevated CO2 and temperature. While cell densities remained constant within each treatment and throughout the experiment, concentrations of HMW-dCCHO, pCCHO and TEP were generally higher under enhanced nutrient stress. At μ = 0.3 d−1, pCCHO concentration increased significantly with elevated CO2 and temperature. At μ = 0.1 d−1, the contribution (mol % C) of HMW-dCCHO to DOC was lower at elevated CO2 and temperature while pCCHO and TEP concentrations were higher. This was most pronounced under greenhouse conditions. Our findings suggest a stronger transformation of primary produced DOC into POC by coagulation of exudates under nutrient limitation. Our results further imply that elevated CO2 and temperature will increase exudation by E. huxleyi and may affect organic carbon partitioning in the ocean due to an enhanced transfer of HMW-dCCHO to TEP by aggregation processes.

Description: Increasing concentrations of dissolved inorganic carbon (DIC) in the interior ocean is expected as a direct consequence of increasing concentrations of CO2 in the atmosphere. This extra DIC is often referred to as anthropogenic carbon (Cant), and its inventory, or increase rate, in the interior ocean has previously been estimated by a multitude of observational approaches. Each of these methods are associated with hard to test assumptions since Cant cannot be directly observed. Results from a simpler concept with few assumptions applied to the Atlantic Ocean are reported on here using two large data collections of carbon relevant bottle data. The change in column inventory on decadal time scales, i.e. the storage rate, of DIC, respiration compensated DIC and oxygen is calculated for the Atlantic Ocean. The average storage rates for DIC and oxygen is calculated to 0.72 ± 1.22 (95% confidence interval of the mean trend: 0.65–0.78) mol m−2 yr−1 and −0.54 ± 1.64 (95% confidence interval of the mean trend: –0.64–(−0.45)) mol m−2 yr−1, respectively, for the Atlantic Ocean, where the uncertainties reflect station-to-station variability and where the mean trends are non-zero at the 95% confidence level. The standard deviation mainly reflects uncertainty due to regional variations, whereas the confidence interval reflects the mean trend. The storage rates are similar to changes found by other studies, although with large uncertainty. For the subpolar North Atlantic the storage rates show significant temporal variation of all variables. This seems to be due to variations in the prevalence of subsurface water masses with different DIC concentrations leading to sometimes different signs of storage rates for DIC and Cant. This study suggest that accurate assessment of the uptake of CO2 by the oceans will require accounting not only for processes that influence Cant but also additional processes that modify CO2 storage.

Description: We conducted measurements of the five important short-lived organic bromine species in the marine boundary layer (MBL). Measurements were made in the Northern Hemisphere mid-latitudes (Sylt Island, North Sea) in June 2009 and in the tropical Western Pacific during the TransBrom ship campaign in October 2009. For the one-week time series on Sylt Island, mean mixing ratios of CHBr3, CH2Br2, CHBr2Cl and CH2BrCl were 2.0, 1.1, 0.2, 0.1 ppt, respectively. We found maxima of 5.8 and 1.6 ppt for the two main components CHBr3 and CH2Br2. Along the cruise track in the Western Pacific (between 41° N and 13° S) we measured mean mixing ratios of 0.9, 0.9, 0.2, 0.1 and 0.1 ppt for CHBr3, CH2Br2, CHBrCl2, CHBr2Cl and CH2BrCl. Air samples with coastal influence showed considerably higher mixing ratios than the samples with open ocean origin. Correlation analyses of the two data sets yielded strong linear relationships between the mixing ratios of four of the five species (except for CH2BrCl). Using a combined data set from the two campaigns and a comparison with the results from two former studies, rough estimates of the molar emission ratios between the correlated substances were: 9/1/0.35/0.35 for CHBr3/CH2Br2/CHBrCl2/CHBr2Cl. Additional measurements were made in the tropical tropopause layer (TTL) above Teresina (Brazil, 5° S) in June 2008, using balloon-borne cryogenic whole air sampling technique. Near the level of zero clear-sky net radiative heating (LZRH) at 14.8 km about 2.25 ppt organic bromine was bound to the five short-lived species, making up 13% of total organic bromine (17.82 ppt). CH2Br2 (1.45 ppt) and CHBr3 (0.56 ppt) accounted for 90% of the budget of short-lived compounds in that region. Near the tropopause (at 17.5 km) organic bromine from these substances was reduced to 1.35 ppt, with 1.07 and 0.12 ppt attributed to CH2Br2 and CHBr3, respectively.

Description: Earth System Climate Models (ESCMs) are valuable tools that can be used to gain a better understanding of the climate system, global biogeochemical cycles and how anthropogenically-driven changes may affect them. Here we describe improvements made to the marine biogeochemical ecosystem component of the University of Victoria's ESCM (version 2.9). Major changes include corrections to the code and equations describing phytoplankton light limitation and zooplankton grazing, the implementation of a more realistic zooplankton growth and grazing model, and the implementation of an iron limitation scheme to constrain phytoplankton growth. The new model is evaluated after a 10 000-yr spin-up and compared to both the previous version and observations. For the majority of biogeochemical tracers and ecosystem processes the new model shows significant improvements when compared to the previous version and evaluated against observations. Many of the improvements are due to better simulation of seasonal changes in higher latitude ecosystems and the effect that this has on ocean biogeochemistry. This improved model is intended to provide a basic new ESCM model component, which can be used as is or expanded upon (i.e., the addition of new tracers), for climate change and biogeochemical cycling research.