ALBERT

Description: Nutrient transfer into the sunlit surface ocean by cyclonic eddies is potentially crucial for sustaining primary productivity in the stratified subtropical gyres. However, the nature of productivity enhancements, including the flow of matter to higher trophic levels and its impact on carbon fluxes, remain poorly resolved. Here, we report a detailed assessment of the biogeochemical response to a cyclonic eddy in the subtropical Northwest Pacific via a combination of ship‐based and autonomous platforms. Primary production was enhanced twofold within the eddy core relative to reference sites outside, whereas phytoplankton biomass even decreased. Pico‐phytoplankton (〈 2 μ m) dominated (〉 80%) both phytoplankton biomass and primary production inside and outside the eddy. The stimulated primary production in the eddy core was accompanied by an approximately twofold increase in mesozooplankton abundance, an approximately threefold increase in particle formation in the deep chlorophyll maximum layer, as well as significantly enhanced surface oceanic CO 2 uptake and net community production. We suggest these observations carry important implications for understanding carbon export in the subtropical ocean and highlight the need to include such subtropical eddy features in ocean carbon budget analyses.

Description: Promoting effects of aluminum addition on chlorophyll biosynthesis and growth of two cultured iron‐limited marine diatoms Linbin Zhou CAS Key Laboratory of Tropical Marine Bio‐resources and Ecology, Guangdong Provincial Key Laboratory of Applied Marine Biology, South China Sea Institute of Oceanology Chinese Academy of Sciences Guangzhou China Marine Biogeochemistry Division GEOMAR Helmholtz Centre for Ocean Research Kiel Germany University of Chinese Academy of Sciences Beijing China https://orcid.org/0000-0001-7230-4116 Fengjie Liu Marine Biogeochemistry Division GEOMAR Helmholtz Centre for Ocean Research Kiel Germany Grantham Institute—Climate Change and the Environment, Department of Life Sciences Imperial College London London UK Eric P. Achterberg Marine Biogeochemistry Division GEOMAR Helmholtz Centre for Ocean Research Kiel Germany Anja Engel Marine Biogeochemistry Division GEOMAR Helmholtz Centre for Ocean Research Kiel Germany https://orcid.org/0000-0002-1042-1955 Peter G.C. Campbell Institut National de la Recherche Scientifique Centre Eau Terre Environnement Quebec Canada https://orcid.org/0000-0001-7160-4571 Claude Fortin Institut National de la Recherche Scientifique Centre Eau Terre Environnement Quebec Canada https://orcid.org/0000-0002-2479-1869 Liangmin Huang CAS Key Laboratory of Tropical Marine Bio‐resources and Ecology, Guangdong Provincial Key Laboratory of Applied Marine Biology, South China Sea Institute of Oceanology Chinese Academy of Sciences Guangzhou China University of Chinese Academy of Sciences Beijing China Yehui Tan CAS Key Laboratory of Tropical Marine Bio‐resources and Ecology, Guangdong Provincial Key Laboratory of Applied Marine Biology, South China Sea Institute of Oceanology Chinese Academy of Sciences Guangzhou China University of Chinese Academy of Sciences Beijing China Abstract Aluminum (Al) may play a role in the ocean's capacity for absorbing atmospheric CO 2 via influencing carbon fixation, export, and sequestration. Aluminum fertilization, especially in iron (Fe)‐limited high‐nutrient, low‐chlorophyll ocean regions, has been proposed as a potential CO 2 removal strategy to mitigate global warming. However, how Al addition would influence the solubility and bioavailability of Fe as well as the physiology of Fe‐limited phytoplankton has not yet been examined. Here, we show that Al addition (20 and 100 nM) had little influence on the Fe solubility in surface seawater and decreased the Fe bio‐uptake by 11–22% in Fe‐limited diatom Thalassiosira weissflogii in Fe‐buffered media. On the other hand, the Al addition significantly increased the rate of chlorophyll biosynthesis by 45–60% for Fe‐limited T. weissflogii and 81–102% for Fe‐limited Thalassiosira pseudonana , as well as their cell size, cellular chlorophyll content, photosynthetic quantum efficiency ( F v / F m ) and growth rate. Under Fe‐sufficient conditions, the Al addition still led to an increased growth rate, though the beneficial effects of Al addition on chlorophyll biosynthesis were no longer apparent. These results suggest that Al may facilitate chlorophyll biosynthesis and benefit the photosynthetic efficiency and growth of Fe‐limited diatoms. We speculate that Al addition may enhance intracellular Fe use efficiency for chlorophyll biosynthesis by facilitating the superoxide‐mediated intracellular reduction of Fe(III) to Fe(II). Our study provides new evidence and support for the iron–aluminum hypothesis.

Description: The simulation of deep-sea conditions in laboratories is technically challenging but necessary for experiments that aim at a deeper understanding of physiological mechanisms or host-symbiont interactions of deep-sea organisms. In a proof-of-concept study, we designed a recirculating system for long-term culture (〉2 yr) of deep-sea mussels Gigantidas childressi (previously Bathymodiolus childressi). Mussels were automatically (and safely) supplied with a maximum stable level of ~60 μmol L−1 methane in seawater using a novel methane–air mixing system. Experimental animals also received daily doses of live microalgae. Condition indices of cultured G. childressi remained high over the years, and low shell growth rates could be detected, too, which is indicative of positive energy budgets. Using stable isotope data, we demonstrate that G. childressi in our culture system gained energy, both, from the digestion of methane-oxidizing endosymbionts and from digesting particulate food (microalgae). Limitations of the system, as well as opportunities for future experimental approaches involving deep-sea mussels, are discussed.

Description: Scientific Significance Statement Millions of predator–prey interactions between deep-diving toothed whales and cephalopods occur daily in the dark deep sea. While predatory whales developed traits to detect and hunt their prey, cephalopods had to expand their anti-predatory strategies specialized for visual predators, to counteract acoustic predators. Since toothed whale-cephalopod interactions have never been directly observed in the deep sea, it remains unknown what selective pressures and traits evolved from this arms race. Combining current knowledge, we formalize four hypotheses and associated research approaches that will guide future investigation on oceanic predator–prey systems. We identify whale echolocation as an unprecedented armament to hunt distant prey and propose that deep-sea squids avoid acoustic predators by (1) reducing their acoustic cross-section through body shape and posture, (2) deep-sea migration, and (3) not schooling. Toothed whale predation emerges as a potential driver of the cephalopod live-fast-die-young strategy—which may now leave cephalopods at competitive advantage under global change.

Description: A sense of kuleana (personal responsibility) in caring for the land and sea. An appreciation for laulima (many hands cooperating). An understanding of aloha 'āina (love of the land). The University of Hawai'i at Manoa hosted the 2023 Ecological Dissertations in Aquatic Sciences (Eco-DAS) program, which fostered each of these intentions by bringing together a team of early career aquatic ecologists for a week of networking and collaborative, interdisciplinary project development

Description: Communities and their functioning are jointly shaped by ecological and evolutionary processes that manifest in diversity shifts of their component species and genotypes. How both processes contribute to community functional change over time is rarely studied. We here repeatedly quantified eco-evolutionary contributions to CO2-driven total abundance and mean cell size changes after short-, mid-, and longer-term (80, 168, and 〉 168 d, respectively) in experimental phytoplankton communities. While the CO2-driven changes in total abundance and mean size in the short- and mid-term could be predominantly attributed to ecological shifts, the relative contribution of evolution increased. Over the longer-term, the CO2-effect and underlying eco-evolutionary changes disappeared, while total abundance increased, and mean size decreased significantly independently of CO2. The latter could be presumably attributed to CO2-independent genotype selection which fed back to species composition. In conclusion, ecological changes largely dominated the regulation of environmentally driven phytoplankton functional shifts at first. However, evolutionary changes gained importance with time, and can ultimately feedback on species composition, and thus must be considered when predicting phytoplankton change.

Description: Methane (CH4) is a climate-relevant trace gas that is emitted from the open and coastal oceans in considerable amounts. However, its distribution in remote oceanic areas is largely unknown. To fill this knowledge gap, dissolved CH4 was measured at nine stations at 75°S in the Ross Sea during austral summer in January 2020. CH4 undersaturation (mean: 82 ± 20%) was found throughout the water column. In subsurface waters, the distribution of CH4 mainly resulted from mixing of water masses and in situ consumption, whereas the CH4 concentrations in the surface mixed layer were mainly driven by air–sea exchange and diapycnal diffusion between the surface and subsurface layers, as well as consumption of CH4. With a mean air–sea CH4 flux density of −0.44 ± 0.34 μmol m−2 d−1, the Ross Sea was a substantial sink for atmospheric CH4 during austral summer, which is in contrast with most oceanic regions, which are known sources.

Description: Water column imaging multibeam echo sounder systems (MBESs) are a promising technology for quantitative estimates of the gas bubble volume flow within large gas seepage areas. Considerable progress has been made in recent years toward applicable calibration methods for MBESs as well as developing inversion models to convert acoustically measured backscattering cross sections to gas bubble volume flow. However, MBESs are still not commonly used for quantitative gas flow assessments. A reason for this is the absence of published processing methods that demonstrate how MBES data can be processed to quantitatively represent bubble streams. Here, we present a novel method (echo grid integration) that allows for assessing the aggregated backscattering cross section of targets within horizontal water layers. This derived value enables quantifying bubble stream gas flow rates using existing acoustic inversion methods. The presented method is based on averaging geo-referenced volume backscattering coefficients onto a high-resolution 3D voxel-grid. The results are multiplied with the voxel volume to represent measurements of the total backscattering cross-section within each voxel cell. Individual gridded values cannot be trusted because the beam pattern effects cause the values of individual targets to “smear” over multiple grid-cells. The true aggregated backscattering cross-section is thus estimated as the integral over the grid-cells affected by this smearing. Numerical simulation of MBES data acquisition over known targets assesses the method's validity and quantify it's uncertainty for different, realistic scenarios. The found low measurement bias (〈 1%), and dispersion (〈 5%) are promising for application in gas flow quantification methods.

Description: Narcomedusae play a key role as top-down regulators in the midwater, the largest and most understudied biome on Earth. Here, we used ecological niche modeling in three-dimensions (3D), ecomorphology, and phylogeny, to answer evolutionary and ecological questions about the widespread narcomedusan genus Solmissus. Our phylogenetic analyses confirmed that Solmissus incisa represents a complex of several cryptic species. Both the different genetic clades and tentacle morphotypes were widespread and often overlapped geographically- the main difference in their distribution and ecological niche being depth. This demonstrated the importance of including the third dimension when modeling the distribution of pelagic species. Contrary to our hypothesis, we found the modeled distribution of the Solmissus genus (n = 1444) and both tentacle morphotypes to be mostly driven by low dissolved oxygen values and a salinity of 34, and slightly by depth and temperature. Solmissus spp. were reproducing all year round, with specimens reproducing in slightly warmer waters (up to 1.25 & DEG;C warmer). Our results suggest that Solmissus spp. will likely come out as climate change winners by expanding their distribution when facing ocean deoxygenation and by increasing their reproduction due to global warming. However, because most available midwater data comes from the northern Pacific, this sampling bias was undoubtedly reflected in the output of our ecological niche models, which should be assessed carefully. Our study illustrated the value of online databases including imagery and videography records, for studying midwater organisms and treating midwater biogeographic regions as 3D spaces.

Description: Microbes in the dark ocean are exposed to hydrostatic pressure increasing with depth. Activity rate measurements and biomass production of dark ocean microbes are, however, almost exclusively performed under atmospheric pressure conditions due to technical constraints of sampling equipment maintaining in situ pressure conditions. To evaluate the microbial activity under in situ hydrostatic pressure, we designed and thoroughly tested an in situ microbial incubator (ISMI). The ISMI allows autonomously collecting and incubating seawater at depth, injection of substrate and fixation of the samples after a preprogramed incubation time. The performance of the ISMI was tested in a high-pressure tank and in several field campaigns under ambient hydrostatic pressure by measuring prokaryotic bulk H-3-leucine incorporation rates. Overall, prokaryotic leucine incorporation rates were lower at in situ pressure conditions than under to depressurized conditions reaching only about 50% of the heterotrophic microbial activity measured under depressurized conditions in bathypelagic waters in the North Atlantic Ocean off the northwestern Iberian Peninsula. Our results show that the ISMI is a valuable tool to reliably determine the metabolic activity of deep-sea microbes at in situ hydrostatic pressure conditions. Hence, we advocate that deep-sea biogeochemical and microbial rate measurements should be performed under in situ pressure conditions to obtain a more realistic view on deep-sea biotic processes.

Description: Global warming, bioinvasions, and parasitism affect single-species performances and species interactions, substantially impacting the structure and stability of marine ecosystems. In light of accelerated global change, the information derived from studies focusing on single species and single drivers is insufficient, calling for a multi-stressor approach under near-natural conditions. We investigated the effects of warming (+3°C) on the performance of a benthic community composed of native and invasive macroalgae, consumers and a trematode parasite in a mesocosm setting. We also assessed the effects of warming and parasitism on the survival and growth of gastropods and mussels and the thermal dependency of trematode performance. Our findings show that warming and grazing by infected gastropods had a large detrimental effect on the invasive macroalga growth. Furthermore, the single and interactive effects of parasitism and warming were detrimental to intermediate host survival and growth, especially to large mussels. Finally, cercarial emergence positively correlated to the natural peaks of summer temperatures, while infection intensity in mussels was higher in larger individuals. Our findings suggest that grazing and warming will be detrimental to the invasive macroalga, favoring the native alga. Moreover, parasitism will enhance grazing, especially in summer, when higher temperatures trigger parasite development. However, parasite-enhanced grazing may be buffered by higher mortality or a shift in the size of infected intermediate hosts under warming. Our findings demonstrate how complex effects of ocean warming can be on food webs and how they can be mediated by parasitism and, as a result, influence native and invasive macroalgae differently.

Description: Estuaries regulate carbon cycling along the land-ocean continuum and thus influence carbon export to the ocean, and global carbon budgets. The Elbe Estuary in Germany has been altered by large anthropogenic perturbations, such as widespread heavy metal pollution, minimally treated wastewater before the 1980s, establishment of wastewater treatment plants after the 1990s, and an overall nutrient and pollutant load reduction in the last three decades. Based on an extensive evaluation of key ecosystem variables, and an analysis of the available inorganic and organic carbon records, this study has identified three ecosystem states in recent history: the polluted (1985–1990), transitional (1991–1996), and recovery (1997–2018) states. The polluted state was characterized by very high dissolved inorganic carbon (DIC) and ammonium concentrations, toxic heavy metal levels, dissolved oxygen undersaturation, and low pH. During the transitional state, heavy metal pollution decreased by 〉 50%, and primary production re-established in spring to summer, with weak seasonality in DIC. Since 1997, during the recovery state, DIC seasonality was driven by primary production, and DIC significantly increased by 〉 23 μmol L−1 yr−1 in the mid to lower estuary, indicating that, along with the improvement in water quality the ecosystem state is still changing. Large anthropogenic perturbations can therefore alter estuarine ecosystems (on the order of decades), as well as induce large and complex biogeochemical shifts and significant changes to carbon cycling.

Description: Plankton is a massive and phylogenetically diverse group of thousands of prokaryotes, protists (unicellular eukaryotic organisms), and metazoans (multicellular eukaryotic organisms; Fig. 1). Plankton functional diversity is at the core of various ecological processes, including productivity, carbon cycling and sequestration, nutrient cycling (Falkowski 2012), interspecies interactions, and food web dynamics and structure (D'Alelio et al. 2016). Through these functions, plankton play a critical role in the health of the coastal and open ocean and provide essential ecosystem services. Yet, at present, our understanding of plankton dynamics is insufficient to project how climate change and other human-driven impacts affect the functional diversity of plankton. That limits our ability to predict how critical ecosystem services will change in the future and develop strategies to adapt to these changes.

Description: Phytoplankton stand at the base of the marine food-web, and play a major role in global carbon cycling. Rising CO2 levels and temperatures are expected to enhance growth and alter carbon:nutrient stoichiometry of marine phytoplankton, with possible consequences for the functioning of marine food-webs and the oceanic carbon pump. To date, however, the consistency of phytoplankton stoichiometric responses remains unclear. We therefore performed a meta-analysis on data from experimental studies on stoichiometric responses of marine phytoplankton to elevated pCO2 and 3–5° warming under nutrient replete and limited conditions. Our results demonstrate that elevated pCO2 increased overall phytoplankton C:N (by 4%) and C:P (by 9%) molar ratios under nutrient replete conditions, as well as phytoplankton growth rates (by 6%). Nutrient limitation amplified the CO2 effect on C:N and C:P ratios, with increases to 27% and 17%, respectively. In contrast to elevated pCO2, warming did not consistently alter phytoplankton elemental composition. This could be attributed to species- and study-specific increases and decreases in stoichiometry in response to warming. While our observed moderate CO2-driven changes in stoichiometry are not likely to drive marked changes in food web functioning, they are in the same order of magnitude as current and projected estimations of oceanic carbon export. Therefore, our results may indicate a stoichiometric compensation mechanism for reduced oceanic carbon export due to declining primary production in the near future

Description: Understanding how marine microbial food webs and their ecosystem functions are changing is crucial for projections of the future ocean. Often, simplified food web models are employed and their solutions are only evaluated against available observations of plankton biomass. With such an approach, it remains unclear how different underlying trophic interactions affect interpretations of plankton dynamics and functioning. Here, we quantitatively compare four hypothetical food webs to data from an existing mesocosm experiment using a refined version of the Minimum Microbial Food Web model. Food web representations range from separated food chains to complex food webs featuring additional trophic links including intraguild predation (IGP). Optimization against observations and taking into account model complexity ensures a fair comparison of the different food webs. Although the different optimized model food webs capture the observations similarly well, projected ecosystem functions differ depending on the underlying food web structure and the presence or absence of IGP. Mesh-like food webs dominated by the microbial loop yield higher recycling and net primary production (NPP) than models dominated by the classical diatom-copepod food chain. A high degree of microzooplankton IGP increases NPP and organic matter recycling, but decreases trophic transfer efficiency (TTE) to copepods. Copepod production, the trophic role of copepods, and TTE are more sensitive to initial biomass changes in chain-like than in complex food webs. Measurements resolving trophic interactions, in particular those quantifying IGP, remain essential to reduce model uncertainty and allow sound conclusions for ecosystem functioning in plankton ecosystems.

Description: In this study, we provide a method to quantify the uncertainty associated with sampling particle size distributions (PSD), using a global compilation of Underwater Vision Profiler observations (UVP, version 5). The UVP provides abundant in situ data of the marine PSD on global scales and has been used for a diversity of applications, but the uncertainty associated with its measurements has not been quantified, including how this uncertainty propagates into derived products of interest. We model UVP sampling uncertainty using Bayesian Poisson statistics and provide formulae for the uncertainty associated with a given sampling volume and observed particle count. We also model PSD observations using a truncated power law to better match the low concentration associated with rare large particles as seen by the UVP. We use the two shape parameters from this statistical model to describe changes in the PSD shape across latitude band, season, and depth. The UVP sampling uncertainty propagates into an uncertainty for modeled carbon flux exceeding 50%. The statistical model is used to extend the size interval used in a PSD-derived carbon flux model, revealing a high sensitivity of the PSD-derived flux model to the inclusion of small particles (80–128 μm). We provide avenues to address additional uncertainties associated with UVP-derived carbon flux calculations.

Description: Physiological sensitivity of cold-water corals to ocean change is far less understood than of tropical corals and very little is known about the impacts of ocean acidification and warming on degradative processes of dead coral framework. In a 13-month laboratory experiment, we examined the interactive effects of gradually increasing temperature and pCO2 levels on survival, growth, and respiration of two prominent color morphotypes (colormorphs) of the framework-forming cold-water coral Lophelia pertusa, as well as bioerosion and dissolution of dead framework. Calcification rates tended to increase with warming, showing temperature optima at ~ 14°C (white colormorph) and 10–12°C (orange colormorph) and decreased with increasing pCO2. Net dissolution occurred at aragonite undersaturation (ΩAr 〈 1) at ~ 1000 μatm pCO2. Under combined warming and acidification, the negative effects of acidification on growth were initially mitigated, but at ~ 1600 μatm dissolution prevailed. Respiration rates increased with warming, more strongly in orange corals, while acidification slightly suppressed respiration. Calcification and respiration rates as well as polyp mortality were consistently higher in orange corals. Mortality increased considerably at 14–15°C in both colormorphs. Bioerosion/dissolution of dead framework was not affected by warming alone but was significantly enhanced by acidification. While live corals may cope with intermediate levels of elevated pCO2 and temperature, long-term impacts beyond levels projected for the end of this century will likely lead to skeletal dissolution and increased mortality. Our findings further suggest that acidification causes accelerated degradation of dead framework even at aragonite saturated conditions, which will eventually compromise the structural integrity of cold-water coral reefs.

Description: Nutrients limiting phytoplankton growth in the ocean are a critical control on ocean productivity and can underpin predicted responses to climate change. The extensive western subtropical North Pacific is assumed to be under strong nitrogen limitation, but this is not well supported by experimental evidence. Here, we report the results of 14 factorial nitrogen–phosphorus–iron addition experiments through the Philippine Sea, which demonstrate a gradient from nitrogen limitation in the north to nitrogen–iron co-limitation in the south. While nitrogen limited sites responded weakly to nutrient supply, co-limited sites bloomed with up to ~60-fold increases in chlorophyll a biomass that was dominated by initially undetectable diatoms. The transition in limiting nutrients and phytoplankton growth capacity was driven by a gradient in deep water nutrient supply, which was undetectable in surface concentration fields. We hypothesize that this large-scale phytoplankton response gradient is both climate sensitive and potentially important for regulating the distribution of predatory fish.

Description: The benthic boundary layer plays a crucial role in the exchange of trace metals between surface sediments and the water column. So far it has been difficult to study dissolved–particulate interactions of trace metals in this highly reactive interface layer due to the lack of suitable sampling methods. We developed a new device, called Benthic Trace Profiler (BTP), which enables simultaneous sampling of near-bottom water and suspended particles in high depth resolution within the first 3 m above the seafloor. The device was tested successfully in the Baltic Sea. The concentrations of several trace metals (Co, Ni, Cu, Zn, and Cd) in the collected bottom waters overlapped with concentrations in water column samples above collected with conventional methods. This observation indicates that the sampling device and method is trace metal clean. The trace metals Fe and Mn showed concentration gradients within the benthic boundary layer indicating an upward diffusive flux. This observation is consistent with a diffusive benthic flux of these trace metals across the sediment–water interface, which was independently verified using pore-water profiles. Suspended particles can be used to study precipitation processes and to determine the carrier phases of trace metals. The BTP fulfilled all the intended requirements as it allowed a simultaneous, uncontaminating and oxygen-free sampling of seawater and suspended particles to gather high-resolution profiles of dissolved and particulate trace metal concentrations above the seafloor. The device closes the gap between water column and sediment sampling and helps researchers to better understand trace metal exchange processes across the ocean's lower boundary.

Description: In recent decades, the increase in terrestrial inputs to freshwater and coastal ecosystems, especially occurring at northern latitudes, has led to a process of water color darkening known as “brownification.” To assess how brownification affects plankton community composition and functioning in northern coastal areas, an in situ mesocosm experiment using a highly colored humic substance to simulate a brownification event was performed in a North Atlantic bay (Hopavågen, Norway) in August 2019. Manual sampling for analyses of nutrient concentrations, phytoplankton pigments and zooplankton abundances was combined with high-frequency (every 15 min) monitoring of key environmental variables to investigate the response of the plankton community in terms of oxygen metabolism and community composition. In response to brownification, the oxygen gross primary production (GPP) and community respiration (R) slowed down significantly, by almost one-third. However, GPP and R both decreased to the same extent; thus, the oxygen metabolic balance was not affected. Moreover, the chlorophyll-a concentration significantly decreased under brownification, by 9% on average, and the chemotaxonomic pigment composition of the phytoplankton changed, indicating their acclimation to the reduced light availability. In addition, brownification seemed to favor appendicularians, the dominant mesozooplankton group in the mesocosms, which potentially contributed to lowering the phytoplankton biomass. In conclusion, the results of this in situ mesocosm experiment suggest that brownification could induce significant changes in phytoplankton and zooplankton community composition and significantly alter the overall oxygen metabolism of plankton communities in a northern Atlantic bay.

Description: In coastal marine environments, physical and biological forces can cause dynamic pH fluctuations from microscale (diffusive boundary layer [DBL]) up to ecosystem‐scale (benthic boundary layer [BBL]). In the face of ocean acidification (OA), such natural pH variations may modulate an organism's response to OA by providing temporal refugia. We investigated the effect of pH fluctuations, generated by the brown alga Fucus serratus' biological activity, on the calcifying epibionts Balanus improvisus and Electra pilosa under OA. For this, both epibionts were grown on inactive and biologically active surfaces and exposed to (1) constant pH scenarios under ambient (pH 8.1) or OA conditions (pH 7.7), or (2) oscillating pH scenarios mimicking BBL conditions at ambient (pH 7.7–8.6) or OA scenarios (pH 7.4–8.2). Furthermore, all treatment combinations were tested at 10°C and 15°C. Against our expectations, OA treatments did not affect epibiont growth under constant or fluctuating (BBL) pH conditions, indicating rather high robustness against predicted OA scenarios. Furthermore, epibiont growth was hampered and not fostered on active surfaces (fluctuating DBL conditions), indicating that fluctuating pH conditions of the DBL with elevated daytime pH do not necessarily provide temporal refugia from OA. In contrast, results indicate that factors other than pH may play larger roles for epibiont growth on macrophytes (e.g., surface characteristics, macrophyte antifouling defense, or dynamics of oxygen and nutrient concentrations). Warming enhanced epibiont growth rates significantly, independently of OA, indicating no synergistic effects of pH treatments and temperature within their natural temperature range.

Description: Lateral fluxes (i.e., outwelling) of dissolved organic (DOC) and inorganic (DIC) carbon and total alkalinity were estimated using radium isotopes at the groundwater, mangrove creek, and continental shelf scales in the Amazon region. Observations of salinity and radium isotopes in the creek indicated tidally driven groundwater exchange as the main source of carbon. Radium-derived transport rates indicate that mangrove carbon is exported out of the continental shelf on timescales of 22 ± 7 d. Bicarbonate was the main form (82% ± 11%) of total dissolved carbon in all samples, followed by DOC (13% ± 12%) and CO2 (5% ± 4%). DIC (18.7 ± 15.7 mmol m−2 d−1) exceeded DOC (3.0 ± 4.1 mmol m−2 d−1) outwelling at all spatial scales. The interpretation of outwelling across the mangrove-ocean continuum is related to the spatial and temporal scales investigated. At all scales, outwelling represented a major coastal carbon pathway driving bicarbonate storage in the ocean.

Description: Global change impacts marine organisms and communities mainly through ocean warming, acidification, deoxygenation, and changes in nutrient inputs and water circulation. To assess the ecological impacts of global change, the effects of multiple interacting environmental drivers, including their fluctuations, should be tested at different levels of biological organization. In an outdoor mesocosm study, we investigated the differential effects of three simulated upwelling events coupled with ocean warming (1–5°C above ambient) on a temperate benthic community in the Western Baltic Sea. Ocean warming, especially in summer when temperatures are close to or above the physiological optimum of many species, is likely to impose thermal stress with species-specific impacts. As the properties of deep water vary seasonally, so will the effects of upwelling. Upwelling of cooler deep water in midsummer may alleviate thermal stress, although this mitigation may be modulated by upwelling-associated shifts in other water-quality parameters such as salinity, nutrients, or late-summer hypoxia. This investigation showed that in the Western Baltic Ocean warming was rather beneficial in early and late summer but detrimental when ambient temperatures were highest in midsummer. The effects of upwelling in the absence of ocean warming were generally weakly beneficial, while this effect tended to vanish with intensifying imposed ocean warming. Hypoxia associated with the late summer upwelling impacted some of the grazer species but did not impact the macroalgae. We conclude that in coastal temperate benthic communities, ocean warming is the predominant stressor that may partially and seasonally be buffered by upwelling.

Description: Fjord ecosystems of the high Arctic are distinct from fjords of temperate latitudes due to the influence of glaciers, icebergs, sea ice, and the permanently low temperatures. The sediment microbiology and biogeochemical processes were analyzed during an international research program with multiple field studies in Svalbard, situated between the Barents Sea and the Arctic Ocean. We here describe the physical and geochemical setting and the predominant microbiological processes in several fjords. Physiological studies of sediments and pure cultures show how the predominantly psychrophilic bacteria are adapted to the near-zero temperature. The microbial communities include bacteria responsible for organic matter hydrolytic degradation, fermentation, and terminal oxidation to CO2. These processes drive the cycling of carbon, sulfur, iron, and manganese. The balance between the dominant sediment microbial processes changes along transects out through the fjords, reflecting the varying impact of the glacier-derived rock flour, rich in metal oxides at the head, and the plankton-derived, labile, marine organic matter at the mouth. Due to accelerated warming of Arctic ecosystems, glaciers are retreating with impacts on the physical, chemical, and biological functioning of the fjord sediment ecosystems.

Description: The Arctic Ocean is highly susceptible to climate change as evidenced by rapid warming and the drastic loss of sea ice during summer. The consequences of these environmental changes for the microbial cycling of organic matter are largely unexplored. Here, we investigated the distribution and composition of dissolved organic matter (DOM) along with heterotrophic bacterial activity in seawater and sea ice of the Eurasian Basin at the time of the record ice minimum in 2012. Bacteria in seawater were highly responsive to fresh organic matter and remineralized on average 55% of primary production in the upper mixed layer. Correlation analysis showed that the accumulation of dissolved combined carbohydrates (DCCHO) and dissolved amino acids (DAA), two major components of fresh organic matter, was related to the drawdown of nitrate. Nitrate‐depleted surface waters at stations adjacent to the Laptev Sea showed about 25% higher concentrations of DAA than stations adjacent to the Barents Sea and in the central Arctic basin. Carbohydrate concentration was the best predictor of heterotrophic bacterial activity in sea ice. In contrast, variability in sea‐ice bacterial biomass was largely driven by differences in ice thickness. This decoupling of bacterial biomass and activity may mitigate the negative effects of biomass loss due to ice melting on heterotrophic bacterial functions. Overall, our results reveal that changes in DOM production and inventories induced by sea‐ice loss have a high potential to enhance the bacterial remineralization of organic matter in seawater and sea ice of the Arctic Ocean.

Description: Sponges are ubiquitous components of various deep‐sea habitats, including cold water coral reefs, and form deep‐sea sponge grounds. Although the deep sea is generally considered to be a food‐limited environment, these ecosystems are known to be hotspots of biodiversity and carbon cycling. To assess the role of sponges in the carbon cycling of deep‐sea ecosystems, we studied the carbon budgets of six dominant deep‐sea sponges of different phylogenetic origin, with various growth forms and hosting distinct associated microbial communities, in an ex situ aquarium setup. Additionally, we determined biomass metrics—planar surface area, volume, wet weight, dry weight (DW), ash‐free dry weight, and organic carbon (C) content—and conversion factors for all species. Oxygen (O2) removal rates averaged 3.3 ± 2.8 μmol O2 g DWsponge h−1 (mean ± SD), live particulate (bacterio‐ and phytoplankton) organic carbon removal rates averaged 0.30 ± 0.39 μmol C g DWsponge h−1 and dissolved organic carbon (DOC) removal rates averaged 18.70 ± 25.02 μmol C g DWsponge h−1. Carbon mass balances were calculated for four species and revealed that the sponges acquired 1.3–6.6 times the amount of carbon needed to sustain their minimal respiratory demands. These results indicate that irrespective of taxonomic class, growth form, and abundance of microbial symbionts, DOC is responsible for over 90% of the total net organic carbon removal of deep‐sea sponges and allows them to sustain themselves in otherwise food‐limited environments on the ocean floor.

Description: Dinitrogen (N2) fixation is a major source of bioavailable nitrogen to oligotrophic ocean communities. Yet, we have limited understanding how ongoing climate change could alter N2 fixation. Most of our understanding is based on short-term laboratory experiments conducted on individual N2-fixing species whereas community-level approaches are rare. In this longer-term in situ mesocosm study, we aimed to improve our understanding on the role of rising atmospheric carbon dioxide (CO2) and simulated deep water upwelling on N2 and carbon (C) fixation rates in a natural oligotrophic plankton community. We deployed nine mesocosms in the subtropical North Atlantic Ocean and enriched seven of these with CO2 to yield a range of treatments (partial pressure of CO2, pCO2 = 352–1025 μatm). We measured rates of N2 and C fixation in both light and dark incubations over the 55-day study period. High pCO2 negatively impacted light and dark N2 fixation rates in the oligotrophic phase before simulated upwelling, while the effect reversed in the light N2 fixation rates in the bloom decay phase after added nutrients were consumed. Dust deposition and simulated upwelling of nutrient-rich deep water increased N2 fixation rates and nifH gene abundances of selected clades including the unicellular diazotrophic cyanobacterium clade UCYN-B. Elevated pCO2 increased C fixation rates in the decay phase. We conclude that elevated pCO2 and pulses of upwelling have pronounced effects on diazotrophy and primary producers, and upwelling and dust deposition modify the pCO2 effect in natural assemblages.

Description: Climate change imposes unusual long‐term trends in environmental conditions, plus some tremendous shifts in short‐term environmental variability, exerting additional stress on marine ecosystems. This paper describes an empirical method that aims to improve our understanding of the performance of benthic filter feeders experiencing changes in environmental conditions, such as temperature, on time scales of minutes to hours, especially during daily cycles or extreme events such as marine heatwaves or hypoxic upwelling. We describe the Fluorometer and Oximeter equipped Flow‐through Setup (FOFS), experimental design, and methodological protocols to evaluate the flood of data, enabling researchers to monitor important energy budget traits, including filtration and respiration of benthic filter‐feeders in response to fine‐tuned environmental variability. FOFS allows online recording of deviations in chlorophyll and dissolved oxygen concentrations induced by the study organism. Transparent data processing through Python scripts provides the possibility to adjust procedures to needs when working in different environmental contexts (e.g., temperature vs. pH, salinity, oxygen, biological cues) and with different filter‐feeding species. We successfully demonstrate the functionality of the method through recording responses of Baltic Sea blue mussels (Mytilus) during one‐day thermal cycles. This method practically provides a tool to help researchers exposing organisms to environmental variability for some weeks or months, to relate the observed long‐term performance responses to short‐term energy budget responses, and to explain their findings with the potential to generalize patterns. The method, therefore, allows a more detailed description of stress‐response relationships and the detection of species' tolerance limits.

Description: The increasing demand for metals is pushing forward the progress of deep‐sea mining industry. The abyss between the Clarion and Clipperton Fracture Zones (CCFZ), a region holding a higher concentration of minerals than land deposits, is the most targeted area for the exploration of polymetallic nodules worldwide, which may likely disturb the seafloor across large areas and over many years. Effects from nodule extraction cause acute biodiversity loss of organisms inhabiting sediments and polymetallic nodules. Attention to deep‐sea ecosystems and their services has to be considered before mining starts but the lack of basic scientific knowledge on the methodologies for the ecological surveys of fauna in the context of deep‐sea mining impacts is still scarce. We review the methodology to sample, process and investigate metazoan infauna both inhabiting sediments and nodules dwelling on these polymetallic‐nodule areas. We suggest effective procedures for sampling designs, devices and methods involving gear types, sediment processing, morphological and genetic identification including metabarcoding and proteomic fingerprinting, the assessment of biomass, functional traits, fatty acids, and stable isotope studies within the CCFZ based on both first‐hand experiences and literature. We recommend multi‐ and boxcorers for the quantitative assessments of meio‐ and macrofauna, respectively. The assessment of biodiversity at species level should be focused and/or the combination of morphological with metabarcoding or proteomic fingerprinting techniques. We highlight that biomass, functional traits, and trophic markers may provide critical insights for biodiversity assessments and how statistical modeling facilitates predicting patterns spatially across point‐source data and is essential for conservation management.

Description: Bacterial sulfate reduction (SR) is often determined by radiotracer techniques using 35S‐labeled sulfate. In environments featuring simultaneous sulfide oxidation, SR can be underestimated due to re‐oxidation of 35S‐sulfide. Recycling of 35S‐tracer is expected to be high in sediment with low concentrations of pore‐water sulfide and high abundance of giant filamentous sulfur‐oxidizing bacteria (GFSOB). Here, we applied a sulfide‐spiking method, originally developed for water samples, to sediments along a shelf‐slope transect (72, 128, 243, 752 m water depth) traversing the Peruvian oxygen minimum zone. Sediment spiked with unlabeled sulfide prior to 35S‐sulfate injection to prevent radiotracer recycling was compared to unspiked sediment. At stations characterized by low natural sulfide and abundant GFSOB (128 and 243 m), the method revealed 1–3 times higher SR rates in spiked sediment. Spiking had no effect on SR in sediment with high natural sulfide despite presence of GFSOB (72 m). Bioturbated sediment devoid of GFSOB (752 m) showed elevated SR in spiked samples, likely from artificial introduction of sulfidic conditions. Sulfide oxidation rates at the 128 and 243 m station, derived from the difference in SR between spiked and unspiked sediment, approximated rates of dissimilatory nitrate reduction to ammonium by GFSOB. Gross SR contributed considerably to benthic dissolved inorganic carbon fluxes at the three shallowest station, confirming that SR is an important process for benthic carbon respirations within the oxygen minimum zone. We recommend to further explore the spiking method to capture SR in sediment featuring low sulfide concentrations and high sulfur cycling by GFSOB.

Description: Prochlorococcus and Synechococcus are the most abundant free-living photosynthetic microorganisms in the ocean. Uncultivated lineages of these picocyanobacteria also thrive in the dimly illuminated upper part of oxygen-deficient zones (ODZs), where an important portion of ocean nitrogen (N) loss takes place via denitrification and anaerobic ammonium oxidation. Recent metagenomic studies revealed that ODZ Prochlorococcus have the genetic potential for using different N forms, including nitrate and nitrite, uncommon N sources for Prochlorococcus, but common for Synechococcus. To determine which N sources ODZ picocyanobacteria are actually using in nature, the cellular N-15 natural abundance (delta N-15) and assimilation rates of different N compounds were determined using cell sorting by flow cytometry and mass spectrometry. The natural delta N-15 of the ODZ Prochlorococcus varied from -4.0 parts per thousand to 13.0 parts per thousand (n = 9), with 50% of the values in the range of -2.1-2.6 parts per thousand. While the highest values suggest nitrate use, most observations indicate the use of nitrite, ammonium, or a mixture of N sources. Meanwhile, incubation experiments revealed potential assimilation rates of ammonium and urea in the same order of magnitude as that expected for total N in several environments including ODZs, whereas rates of nitrite and nitrate assimilation were very low. Our results thus indicate that reduced forms of N and nitrite are the dominant sources for ODZ picocyanobacteria, although nitrate might be important on some occasions. ODZ picocyanobacteria might thus represent potential competitors with anammox bacteria for ammonium and nitrite, with ammonia-oxidizing archaea for ammonium, and with nitrite-oxidizing bacteria for nitrite.

Description: The plea for using more “realistic,” community‐level, investigations to assess the ecological impacts of global change has recently intensified. Such experiments are typically more complex, longer, more expensive, and harder to interpret than simple organism‐level benchtop experiments. Are they worth the extra effort? Using outdoor mesocosms, we investigated the effects of ocean warming (OW) and acidification (OA), their combination (OAW), and their natural fluctuations on coastal communities of the western Baltic Sea during all four seasons. These communities are dominated by the perennial and canopy‐forming macrophyte Fucus vesiculosus—an important ecosystem engineer Baltic‐wide. We, additionally, assessed the direct response of organisms to temperature and pH in benchtop experiments, and examined how well organism‐level responses can predict community‐level responses to the dominant driver, OW. OW affected the mesocosm communities substantially stronger than acidification. OW provoked structural and functional shifts in the community that differed in strength and direction among seasons. The organism‐level response to OW matched well the community‐level response of a given species only under warm and cold thermal stress, that is, in summer and winter. In other seasons, shifts in biotic interactions masked the direct OW effects. The combination of direct OW effects and OW‐driven shifts of biotic interactions is likely to jeopardize the future of the habitat‐forming macroalga F. vesiculosus in the Baltic Sea. Furthermore, we conclude that seasonal mesocosm experiments are essential for our understanding of global change impact because they take into account the important fluctuations of abiotic and biotic pressures.

Description: The duration and magnitude of the North Atlantic spring bloom impacts both higher trophic levels and oceanic carbon sequestration. Nutrient exhaustion offers a general explanation for bloom termination, but detail on which nutrients and their relative influence on phytoplankton productivity, community structure, and physiology is lacking. Here, we address this using nutrient addition bioassay experiments conducted across the midlatitude North Atlantic in June 2017 (late spring). In four out of six experiments, phytoplankton accumulated over 48–72 h following individual additions of either iron (Fe) or nitrogen (N). In the remaining two experiments, Fe and N were serially limiting, that is, their combined addition sequentially enhanced phytoplankton accumulation. Silicic acid (Si) added in combination with N + Fe led to further chlorophyll a (Chl a) enhancement at two sites. Conversely, addition of zinc, manganese, cobalt, vitamin B12, or phosphate in combination with N + Fe did not. At two sites, the simultaneous supply of all six nutrients, in combination with N + Fe, also led to no further Chl a enhancement, but did result in an additional 30–60% particulate carbon accumulation. This particulate carbon accumulation was not matched by a Redfield equivalent of particulate N, characteristic of high C:N organic exudates that enhance cell aggregation and sinking. Our results suggest that growth rates of larger phytoplankton were primarily limited by Fe and/or N, making the availability of these nutrients the main bottom‐up factors contributing to spring bloom termination. In addition, the simultaneous availability of other nutrients could modify bloom characteristics and carbon export efficiency.

Description: Numerical models are a suitable tool to quantify impacts of predicted climate change on complex ecosystems but are rarely used to study effects on benthic macroalgal communities. Fucus vesiculosus L. is a habitat-forming macroalga in the Baltic Sea and alarming shifts from the perennial Fucus community to annual filamentous algae are reported. We developed a box model able to simulate the seasonal growth of the Baltic Fucus-grazer-epiphyte system. This required the implementation of two state variables for Fucus biomass in units of carbon (C) and nitrogen (N). Model equations describe relevant physiological and ecological processes, such as storage of C and N assimilates by Fucus, shading effects of epiphytes or grazing by herbivores on both Fucus and epiphytes, but with species-specific rates and preferences. Parametrizations of the model equations and required initial conditions were based on measured parameters and process rates in the near-natural Kiel Outdoor Benthocosm (KOB) experiments during the Biological Impacts of Ocean Acidification project. To validate the model, we compared simulation results with observations in the KOB experiment that lasted from April 2013 until March 2014 under ambient and climate-change scenarios, that is, increased atmospheric temperature and partial pressure of carbon dioxide. The model reproduced the magnitude and seasonal cycles of Fucus growth and other processes in the KOBs over 1 yr under different scenarios. Now having established the Fucus model, it will be possible to better highlight the actual threat of climate change to the Fucus community in the shallow nearshore waters of the Baltic Sea.

Description: Numerical simulations of ocean biogeochemical cycles need to adequately represent particle sinking velocities (SV). For decades, Stokes' Law estimating particle SV from density and size has been widely used. But while Stokes' Law holds for small, smooth, and rigid spheres settling at low Reynolds number, it fails when applied to marine aggregates complex in shape, structure, and composition. Minerals and zooplankton can alter phytoplankton aggregates in ways that change their SV, potentially improving the applicability of Stokes' models. Using rolling cylinders, we experimentally produced diatom aggregates in the presence and absence of minerals and/or microzooplankton. Minerals and to a lesser extent microzooplankton decreased aggregate size and roughness and increased their sphericity and compactness. Stokes' Law parameterized with a fractal porosity modeled adequately size‐SV relationships for mineral‐loaded aggregates. Phytoplankton‐only aggregates and those exposed to microzooplankton followed the general Navier‐Stokes drag equation suggesting an indiscernible effect of microzooplankton and a drag coefficient too complex to be calculated with a Stokes' assumption. We compared our results with a larger data set of ballasted and nonballasted marine aggregates. This confirmed that the size‐SV relationships for ballasted aggregates can be simulated by Stokes' models with an adequate fractal porosity parameterization. Given the importance of mineral ballasting in the ocean, our findings could ease biogeochemical model parameterization for a significant pool of particles in the ocean and especially in the mesopelagic zone where the particulate organic matter : mineral ratio decreases. Our results also reinforce the importance of accounting for porosity as a decisive predictor of marine aggregate SV.

Description: Ocean surface partial pressure of carbon dioxide (pCO2) is a key factor controlling air–sea CO2 fluxes. Most surface pCO2 data are collected with relatively large and complex air–water equilibrators coupled to stand‐alone infrared analyzers installed on Ships of OPportunity (SOOP‐CO2). This approach has proven itself through years of successful deployments, but expansion and sustainability of the future measurement network faces challenges in terms of certification, autonomy, and maintenance, which motivates development of new systems. Here, we compare performance of three underway pCO2 measurement systems (General Oceanics, SubCtech, and Pro‐Oceanus), including a recently developed compact flow‐through, sensor‐based system. The systems were intercompared over a period of 34 days during two crossings of the subpolar North Atlantic Ocean. With a mean difference from the General Oceanics system of −5.7 ± 4.0 μatm (Pro‐Oceanus) and −4.7 ± 2.9 μatm (SubCtech) during the 1st crossing, our results indicate potential for good agreement between the systems. The study highlighted the challenge of assuring accuracy over long periods of time, particularly seen in a worse agreement during the 2nd crossing, and revealed a number of sources of systematic errors. These can influence accuracy of the measurements, agreement between systems and include slow response of membrane‐based systems to pCO2 changes, “within‐ship” respiration due to biofouling, and bias in measurement of the temperature of equilibration. These error sources can be controlled or corrected for, however, if unidentified, their magnitude can be significant relative to accuracy criteria assigned to the highest‐quality data in global databases. The advantages of the compact flow‐through system are presented along with a discussion of future solutions for improving data quality.

Description: Eutrophication and climate change will affect habitats of species and more generally, the structure and functioning of ecosystems. We used a three‐dimensional, coupled hydrodynamic‐biogeochemical model to investigate potential future changes in size and location of potential habitats of marine species during the 21st century in a large, eutrophicated brackish sea (the Baltic Sea, northern Europe). We conducted scenario projections under the combined impact of nutrient load and climate change. Possible future changes of the eutrophication state of this sea were also assessed through two policy‐relevant indicators. The results imply a physiologically more stressful environment for marine species by the end of the 21st century: volumes of higher salinity water become more hypoxic/anoxic and the volumes of low salinity, oxic water increase. For example, these results impact and reduce cod reproductive habitats. The decrease is mainly climate change induced in the Baltic basins less directly influenced by inflows of saline, oxic water to the Baltic Sea (E Gotland and Gdansk Basins). In basins more directly influenced by such inflows (Arkona and Bornholm Basins), the combined effect from climate change and nutrient loads is of importance. The results for the eutrophication state indicators clearly indicate a more eutrophic sea than at present without a rigorous nutrient reduction policy, that is, the necessity to implement the Baltic Sea Action Plan. The multidisciplinary, multiscenario assessment strategy presented here provides a useful concept for the evaluation of impacts from cumulative stresses of changing climate and socioeconomic pressures on future eutrophication indicators and habitats of marine species.

Description: Enhanced nutrient input and warming have led to the development of low oxygen (hypoxia) in coastal waters globally. For many coastal areas, insight into redox conditions prior to human impact is lacking. Here, we reconstructed bottom water redox conditions and sea surface temperatures (SSTs) for the coastal Stockholm Archipelago over the past 3000 yr. Elevated sedimentary concentrations of molybdenum indicate (seasonal) hypoxia between 1000b.c.e.and 1500c.e. Biomarker-based (TEX86) SST reconstructions indicate that the recovery from hypoxia after 1500c.e.coincided with a period of significant cooling (similar to 2 degrees C), while human activity in the study area, deduced from trends in sedimentary lead and existing paleobotanical and archeological records, had significantly increased. A strong increase in sedimentary lead and zinc, related to more intense human activity in the 18(th)and 19(th)century, and the onset of modern warming precede the return of hypoxia in the Stockholm Archipelago. We conclude that climatic cooling played an important role in the recovery from natural hypoxia after 1500c.e., but that eutrophication and warming, related to modern human activity, led to the return of hypoxia in the 20(th)century. Our findings imply that ongoing global warming may exacerbate hypoxia in the coastal zone of the Baltic Sea.

Description: While there is a general sense that lakes can act as sentinels of climate change, their efficacy has not been thoroughly analyzed. We identified the key response variables within a lake that act as indicators of the effects of climate change on both the lake and the catchment. These variables reflect a wide range of physical, chemical, and biological responses to climate. However, the efficacy of the different indicators is affected by regional response to climate change, characteristics of the catchment, and lake mixing regimes. Thus, particular indicators or combinations of indicators are more effective for different lake types and geographic regions. The extraction of climate signals can be further complicated by the influence of other environmental changes, such as eutrophication or acidification, and the equivalent reverse phenomena, in addition to other land-use influences. In many cases, however, confounding factors can be addressed through analytical tools such as detrending or filtering. Lakes are effective sentinels for climate change because they are sensitive to climate, respond rapidly to change, and integrate information about changes in the catchment.

Description: Life history responses are expected to accompany climate warming, yet little is known how long-term effects of climate and environmental change affect the seasonal dynamics of planktonic organisms. We used an historical data set from Lake Washington (U.S.A.) to quantify population responses of a calanoid copepod (Leptodiaptomus ashlandi) to long-term changes in temperature and resource availability and explore potential mechanisms for the responses. Increasing water temperatures (annual mean increase of 1.5 degrees C in the upper 10-m water volume) and longer stratification periods (about 4 weeks) were observed between 1962 and 2005, coincident with a pronounced decline in Leptodiaptomus densities. However, production was maintained because of an increase in the production to biomass ratio and a life cycle shift in Leptodiaptomus from an annual to a 6-month cycle. Cross-wavelet analyses demonstrated that the annual thermal forcing of copepod recruitment observed during the first two decades of the study weakened substantially, leading to more stochastic population dynamics during the past two decades. This shift from one to two generations per year was most likely produced by a longer and warmer growing period combined with changing fluctuations in resource (phytoplankton) availability. Climate change can lead to higher-frequency voltinism in ectothermic organisms and to temporal reorganization of their population dynamics.

Description: Thousands of moribund thaliacean carcasses (Pyrosoma atlanticum) were deposited between February and March 2006 at the seafloor in the Ivory Coast area (West Africa). Remotely operated vehicle surveys were conducted in a continuous depth gradient between 20 and 1275 m along an oil pipeline. Video and still photography were used to estimate the carcass distribution, density, and size on the seabed, as well as recording the local megafauna interactions with the gelatinous material. Large patches of dead pyrosomids covered extensive areas on the continental slope, whereas minor aggregations were found on the shelf. The carcasses were in many instances trapped along the pipelines, accumulating extensively in troughs and furrows in the slope, and especially in soft sediment channels. Pyrosoma atlanticum dried samples were used to calculate the carbon content, enabling the extrapolation to the densities and sizes recorded in the video surveys. The average standing stock of organic carbon associated with the carcasses was 〉5 g C m-2 in the whole slope and canyon, with values as high as 22 g C m-2 in certain areas. Eight megafaunal species from three different phyla were observed 63 times directly feeding on the decomposing carcasses. The gelatinous carbon may have contributed substantially to the detrital food web including microbes at the seabed, and certainly to the diet of larger benthic organisms. The organism-level carbon measurements and documented fate of pyrosomid organic carbon is new evidence of the importance of gelatinous material in large-scale processes and elemental cycling.

Description: To examine the grazing effects of copepod-dominated mesozooplankton on heterotrophic microbial communities, four mesocosm experiments using gradients of zooplankton abundance were carried out at a coastal marine site. The responses of different protist groups (nanoflagellates, ciliates) and bacterioplankton in terms of abundance and additionally, for bacteria, diversity, production, and exoenzymatic activity, were monitored during 1 week of incubation. Independent of the initial experimental abiotic conditions and the dominating copepod species, zooplankton caused order-of-magnitude changes in microbial functional groups in a clear community-wide four-link trophic cascade. The strongest predatory effects were observed for protist concentrations, thus generating inverse relationships between mesozooplankton and ciliates and between ciliates and nanoplankton. Copepod grazing effects propagated even further, not only reducing the abundance, production, and hydrolytic activity of bacterioplankton but also increasing bacterial diversity. The overall strength of this trophic cascade was dampened with respect to bacterial numbers, but more pronounced with respect to bacterial diversity and activity. High predation pressure by heterotrophic nanoflagellates, realized at the highest copepod abundance, was probably the underlying mechanism for these structural changes in the bacterial assemblages. Our results thus suggest a mechanism by which changes in higher trophic levels of marine plankton indirectly affect prokaryotic assemblages and microbially mediated ecosystem functions.

Description: We examined the physiological responses of steady-state iron (Fe)-replete and Fe-limited cultures of the biogeochemically critical marine unicellular diazotrophic cyanobacterium Crocosphaera at glacial (19 Pa; 190 ppm), current (39 Pa; 380 ppm), and projected year 2100 (76 Pa; 750 ppm) CO2 levels. Rates of N2 and CO2 fixation and growth increased in step with increasing partial pressure of CO2 (pCO2), but only under Fe-replete conditions. N2 and carbon fixation rates at 75 Pa CO2 were 1.4-1.8-fold and 1.2-2.0-fold higher, respectively, relative to those at present day and glacial pCO2 levels. In Fe-replete cultures, cellular Fe and molybdenum quotas varied threefold and were linearly related to N2 fixation rates and to external pCO2. However, N2 fixation and trace metal quotas were decoupled from pCO2 in Fe-limited Crocosphaera. Higher CO2 and Fe concentrations both resulted in increased cellular pigment contents and affected photosynthesis vs. irradiance parameters. If these results also apply to natural Crocosphaera populations, anthropogenic CO2 enrichment could substantially increase global oceanic N2 and CO2 fixation, but this effect may be tempered by Fe availability. Possible biogeochemical consequences may include elevated inputs of new nitrogen to the ocean and increased potential for Fe and/or phosphorus limitation in the future high-CO2 ocean, and feedbacks to atmospheric pCO2 in both the near future and over glacial to interglacial timescales.

Description: Nutrient addition bioassay experiments were performed in the low-nutrient, low-chlorophyll oligotrophic subtropical North Atlantic Ocean to investigate the influence of nitrogen (N), phosphorus (P), and/or iron (Fe) on phytoplankton physiology and the limitation of primary productivity or picophytoplankton biomass. Additions of N alone resulted in 1.5-2 fold increases in primary productivity and chlorophyll after 48 h, with larger (~threefold) increases observed for the addition of P in combination with N (NP). Measurements of cellular chlorophyll contents permitted evaluation of the physiological response of the photosynthetic apparatus to N and P additions in three picophytoplankton groups. In both Prochlorococcus and the picoeukaryotes, cellular chlorophyll increased by similar amounts in N and NP treatments relative to all other treatments, suggesting that pigment synthesis was N limited. In contrast, the increase of cellular chlorophyll was greater in NP than in N treatments in Synechococcus, suggestive of NP co-limitation. Relative increases in cellular nucleic acid were also only observed in Synechococcus for NP treatments, indicating co-limitation of net nucleic acid synthesis. A lack of response to relief of nutrient stress for the efficiency of photosystem II photochemistry, Fv :Fm, suggests that the low nutrient supply to this region resulted in a condition of balanced nutrient limited growth, rather than starvation. N thus appears to be the proximal (i.e. direct physiological) limiting nutrient in the oligotrophic sub-tropical North Atlantic. In addition, some major picophytoplankton groups, as well as overall autotrophic community biomass, appears to be co-limited by N and P.

Description: We investigated potential connections between climate and the population size of gelatinous carnivores in the northwestern Mediterranean. The interannual variability of large- and meso-scale climate factors was related to gelatinous carnivore dynamics by means of a multivariate regression model. Gelatinous carnivore population dynamics integrate the climate-related changes in the northwestern Mediterranean with close correlations between climate and the population size of the Calycophoran siphonophores Chelophyes appendiculata and Abylopsis tetragona and the Hydromedusae Rhopalonema velatum and Solmundella bitentaculata. We show that the sensitivity of particular gelatinous carnivore species to environmental forcing relates to their seasonal appearance and peak of abundance. Our results advocate the occurrence of short time windows, during which gelatinous carnivores appear more sensitive to environmental conditions, and favorable conditions may therefore substantially enhance their annual peaks. Furthermore, there were threshold values from which climate effects on gelatinous carnivores become noticeable, indicating that the climate-gelatinous carnivore relationship intensifies according to the strength of climate forcing. The possibility of using the North Atlantic climate variability for assessing and predicting interannual abundance changes of these organisms in the northwestern Mediterranean is considered.

Description: Identification of the proximal nutrient limiting primary production is a necessary first step toward evaluating the physiological state of phytoplankton communities and the biogeochemical constraints on the current oceanic carbon cycle. We conducted 48-h nutrient addition bioassay experiments to evaluate nitrogen, phosphorus, and iron limitation of primary productivity, net chlorophyll synthesis, and net increase in cell numbers of the dominant picophytoplankton from the tropical North Atlantic. Our results indicate that N was the proximal limiting factor for primary production during the autumn of 2002, followed by P and then Fe. Net chlorophyll synthesis was significantly stimulated by addition of N alone and further stimulated by addition of P. Analysis of picophytoplankton populations by analytical flow cytometry revealed a more complex response. Cellular red fluorescence, an index of cell chlorophyll content, increased in Prochlorococcus, Synechococcus, and picoeukaryotes in response to addition of NH4NO3 but was not affected by single or combined additions of P and Fe. In contrast, cell abundance in these picophytoplankton populations increased only after combined N and P (63% of comparisons) or N and Fe (21% of comparisons) additions. Thus, our experiments revealed that chlorophyll synthesis and primary production were limited by the availability of nitrogen alone, while net increase in cell abundance was colimited by N and P or N and Fe in the majority of these picophytoplankton populations.

Description: Bacterial productivity and biomass are thought to be limited by dissolved organic carbon (DOC) in much of the world’s oceans. However, the mixed layer of oligotrophic oceans is often depleted in dissolved inorganic nitrogen and phosphate, raising the possibility that macronutrients may also limit heterotrophic bacterial growth. We used nutrient bioassay experiments to determine whether inorganic nutrients (N, P, Fe) and/or DOC could limit bacterial productivity and biomass in the central North Atlantic during the spring of 2004 (Mar–Apr). We observed that both heterotrophic bacterial productivity and biomass were co-limited by N and P in the oligotrophic North Atlantic, and additions of labile DOC (glucose) provided no stimulation unless N and P were also added. Flow cytometry results indicated that only a small subset of large cells high in nucleic acid content were responsible for the increased productivity in the combined NP amendments. In contrast, nutrient additions elicited no net change on the dominant component of the bacterial population, composed of small cells with relatively low nucleic acid content. In the combined NP treatments the relative increase in bacterial production was greater than that measured when phytoplankton productivity was relieved of nitrogen limitation. These results suggest that N and P co-limitation in the bacterial community results in increased competition between the heterotrophic and autotrophic components of the surface communities in the Central North Atlantic Ocean, and potentially impacts the cycling of organic matter by the bacterioplankton.

Description: Calcification of the cosmopolitan coccolithophore species Emiliania huxleyi was investigated in relation to the cell division cycle with the use of batch cultures. With a 12 : 12 h light : dark cycle, the population was synchronised to undergo division as a cohort, simultaneously passing through the G1 (assimilation), S (DNA replication), and G2+M (cell division and mitosis) phases. Cell division was followed with the use of quantitative DNA staining and flow cytometry. Simultaneously, carbon-14 (14C) assimilation in organic and inorganic carbon as well as cell abundance, size, and organic nitrogen content were measured at 2-h intervals. In additional experiments, changes in calcification and cell cycle stages were investigated in nitrogen-, phosphorus-, and light-limited cultures. Calcification occurred only during the G1 cell cycle phase, as seen by the very tight correlation between the percentage of cells in G1 and calcification during the dark period. When growth was limited by nitrogen, cells decreased in size, remained in the G1 phase, and showed a moderate increase in the cell-specific calcite content. Limitation of growth by phosphorus, however, caused a significant increase in cell size and a dramatic increase in cellular calcite. Light limitation, by slowing the growth rate, prolonged the time cells spent in the G1 phase with a corresponding increase in the cellular calcite content. These results help explain the differing responses of coccolithophorid growth to nitrogen, phosphorus, and light limitation.

Description: Sea ice brines were collected from a single floe composed of different ice types in the western Weddell Sea in December 2004. The chemical composition of the brines (temperature: 23.4°C to 22.1°C; salinity: 40–63) was examined on seven occasions over 25 days with measurements of dissolved oxygen, dissolved inorganic macronutrients (nitrate plus nitrite, ammonium, phosphorus [DIP], and silicic acid), pH, total alkalinity (AT), dissolved organic carbon (DOC) and nitrogen (DON), total dissolved inorganic carbon (CT), and the stable isotopic composition of CT (δ13CT). The in situ pH ranged from 8.41-8.82 on the seawater scale, dissolved oxygen from 212-604 µmol kg−1, nitrate from 0.1-3.1 µmol kg−1, ammonium 0.1-2.4 µmol kg−1, DIP 0.4- 2.0 µmol kg−1, silicic acid 4-80 µmol kg−1, AT 2,690-4,620 µeq kg−1, DOC 115-359 µmol kg−1, DON 8-26 µmol kg−1, CT 2,090-3,550 µmol kg−1, and δ13CT +2.9‰ - +6.4‰. Compared with the chemical composition of surface oceanic water (salinity of 34), the brines had elevated pH, reduced concentrations of dissolved inorganic macronutrients (including carbon), especially dissolved inorganic nitrogen, and were mostly supersaturated with dissolved oxygen with respect to equilibrium with air, whereas the CT was considerably enriched in 13C. The chemical composition of the brines was consistent with internal biological productivity, but there was a lack of a distinctive and uniform relationship among the major dissolved inorganic nutrients typically used for describing biological activity. This was interpreted as the result of varying stoichiometry of biological activity within a very small spatial scale. Modification by abiotic processes was a potential contributing factor, such as degassing acting on the dissolved oxygen concentration. Carbonate mineral formation, acting on AT and CT, was not evident in brines from first-year ice but was apparent in brine from second-year ice.

Description: We investigated whether nutrient limitations of primary producers act upward through food webs only in terms of density effects or if there is a second pathway for nutrient limitation signals channelled upward to higher trophic levels. We used tritrophic food chains to assess the effects of nutrient-limited phytoplankters (the cryptophyte Rhodomonas salina) on herbivorous zooplankters (the calanoid copepod Acartia tonsa) and finally zooplanktivores (larval herring Clupea harengus) living on the herbivores. The primary producers� food quality had a significant effect on fish condition. Our experimental phosphorus-limited food chain resulted in larval fish with a significantly poorer condition than their counterparts reared under nitrogen-limited or nutrient-sufficient conditions. Our results show that mineral nutrient requirements of consumers have to be satisfied first before fatty acids can promote further growth. This challenges the match/mismatch hypothesis, which links larval fish survival probability solely to prey availability, and could imply that reduced nutrient releases into the environment may affect fish stocks even more severely than previously believed.

Description: To investigate the biogeochemistry of iron in the waters of the European continental margin, we determined the dissolved iron distribution and redox speciation in filtered (〈0.2 μm) open-ocean and shelf waters. Depth profiles were sampled over the shelf slope southeast of the Chapelle Bank area (47.61°N, 4.24°W to 46.00°N, 8.01°W) and a horizontal surface-water transect over the shelf and through the English Channel (la Manche) and the southern North Sea (46°N, 8°W to 52°N, 4°E). An abrupt trace-metal front was found near the shelf slope, indicated by a horizontal gradient of dissolved iron (DFe) and aluminium (DAl), which correlated with changing salinities (r2 = 0.572 and 0.528, respectively, n = 92). Labile Fe(II) concentrations varied from 〈12 pmol L-1 in North Atlantic surface waters to >200 pmol L-1 in the near bottom waters of the shelf break. Labile Fe(II) accounted for ∼5 of the dissolved iron species in surface shelf waters (mean 5.0 ± 2.7), whereas higher Fe(II) fractions (i.e., >8) were observed near the sea bottom on the shelf break and during a midday solar maximum in surface waters in the vicinity of the Scheldt river plume. Benthic processes (resuspension and diagenesis) constituted important sources of Fe(II) and DFe in this region, and photoreduction of Fe(III) species in shelf waters caused enhanced labile Fe(II) concentrations. These processes increased the lability of iron and its potential availability to marine organisms in the shelf ecosystem. © 2007, by the American Society of Limnology and Oceanography, Inc.

Description: To date no study exists that directly addresses changes in dynamics of heterotrophic bacteria in surface waters in relation to partial pressure of CO2 (pCO2). Therefore, we studied the effect of changes in pCO2 on bacterial abundance and activities by using mesocosms with different pCO2 levels (~190, ~370, and ~700 ppmV, representing past, present-day, and future atmospheric pCO2, respectively). Abundance of total bacteria did not differ with increasing pCO2 throughout the whole study period, whereas bacterial protein production (BPP) was highest at highest pCO2. This effect was even more pronounced for cell-specific production rates, especially those of attached bacteria, which were up to 25 times higher than those of free bacteria. During the breakdown of the bloom, however, the abundance of both free and attached bacteria was significantly increased with pCO2. Differences in bacterial growth rate (µ) were smaller than those of BPP, but both µ and BPP of attached bacteria were elevated under high pCO2. Averages of total protease as well as α- and α -glucosidase activities were highest at elevated pCO2 levels, but a statistically significant dependence on pCO2 was only evident for protease activity. There is a measurable but indirect effect of changes in pCO2 on bacterial activities that are mainly linked to phytoplankton and presumably particle dynamics

Description: During the European Iron Fertilisation Experiment (EIFEX), performed in the Southern Ocean, we investigated the reactions of different phytoplankton size classes to iron fertilization, applying measurements of size fractionated pigments, particulate organic matter, microscopy, and flow cytometry. Chlorophyll a (Chl a) concentrations at 20-m depth increased more than fivefold following fertilization through day 26, while concentrations of particulate organic carbon (POC), nitrogen (PON), and phosphorus (POP) roughly doubled through day 29. Concentrations of Chl a and particulate organic matter decreased toward the end of the experiment, indicating the demise of the iron-induced phytoplankton bloom. Despite a decrease in total diatom biomass at the end of the experiment, biogenic particulate silicate (bPSi) concentrations increased steadily due to a relative increase of heavily silicified diatoms. Although diatoms 〉 20 mu m were the main beneficiaries of iron fertilization, the growth of small diatoms (2-8 mu m) was also enhanced, leading to a shift from a haptophyte- to a diatom-dominated community in this size fraction. The total biomass had lower than Redfield C : N, N : P, and C : P ratios but did not show significant trends after iron fertilization. This concealed various alterations in the elemental composition of the different size fractions. The microplankton 20 mu m) showed decreasing C : N and increasing N : P and C : P ratios, possibly caused by increased N uptake and the consumption of cellular P pools. The nanoplankton (2-20 mu m) showed almost constant C : N and decreasing N : P and C : P ratios. Our results suggest that the latter is caused by a shift in composition of taxonomic groups.

Description: We quantify, compare, and generalize responses of experimental nutrient loadings (LN) on planktonic community structure and function in coastal waters. Data were derived from three mesocosm experiments undertaken in Baltic (BAL), Mediterranean (MED), and Norwegian (NOR) coastal waters. A planktonic model with seven functional compartments and 30-32 different carbon flows fit to all three experiments was used as a framework for flow-rate estimation and comparison. Flows were estimated on the basis of time series of measured biomass, some measured flows, and inverse modeling. Biomass and gross uptake rate of carbon of most groups increased linearly with increasing LN in the nutrient input range of 0-1 µmol N L-1 d-1 at all locations. The fate of the gross primary production (GPP) was similar in all systems. Autotrophic biomass varied by two orders of magnitude among locations, with the lowest biomass and response to nutrient addition in MED waters. The variation of GPP among sites was less than one order of magnitude. Mesozooplankton dominated by doliolids (Tunicata), but not those dominated by copepods, presumably exerted efficient control of the autotrophic biomass, thereby buffering responses of autotrophs to high nutrient input. Among the many factors that can modify the responses of autotrophs to nutrients, the time scale over which the enrichment is made and the precise mode of nutrient enrichment are important. We suggest a general concept that may contribute to a scientific basis for understanding and managing coastal eutrophication

Description: We investigated the carbon acquisition of three marine microalgae, Skeletonema costatum, Phaeocystis globosa, and Emiliania huxleyi in response to different light regimes. Rates of photosynthetic O2 evolution and CO2 and HCO3- uptake were measured by membrane inlet mass spectrometry in cells acclimated to cycles of 16 : 8 light : dark (LD; h : h) and 12 : 12 LD and were compared with those obtained under continuous light. In addition, cellular leakage was estimated for different photoperiods and ambient CO2 concentrations during growth. Maximum rates of photosynthesis more or less doubled under LD cycles compared with continuous light. In S. costatum and E. huxleyi, a remarkably higher contribution of HCO3- to the overall carbon uptake was observed under LD cycles. In contrast, P. globosa did not change its CO2 :HCO3- uptake ratio in response to daylength. Half saturation concentrations (K1/2) for O2 evolution and inorganic carbon (Ci) uptake were also influenced by the photoperiod. Under LD cycles K1/2 values for photosynthesis in S. costatum and P. globosa were similar or higher compared with continuous light, whereas they were much lower in E. huxleyi. With the exception of CO2 uptake in E. huxleyi and P. globosa, affinities for Ci decreased under the LD cycles. Cellular leakage was highest for E. huxleyi and lowest for S. costatum and generally decreased with increasing CO2 concentration. Although this study confirms species specific differences in the CO2-concentrating mechanisms (CCMs), the effect of daylength on CO2 and HCO3- uptake has hitherto not been described. We put forward the idea that variations in light condition influence the cellular carbon demand, thereby imposing a stronger control on CCM regulation than the naturally occurring changes in CO2 supply.

Description: In recent studies the Sr/Ca ratio of coccolithophore calcite was used as a proxy for past coccolithophore growth and calcification rates. Since Sr and Ca concentrations in seawater have not remained constant through time, interpretation of Sr/Ca data from the coccolith-dominated sedimentary record requires knowledge about the incorporation of seawater Sr into coccolith calcite during coccolithogenesis. Here we show that Sr/Ca of Emiliania huxleyi coccoliths is linearly related to seawater Sr/Ca, meaning that the Sr exchange coefficient does not change with changing seawater Sr/Ca. The exchange coefficient for Sr in this study, 0.39, is close to values presented in the literature and is high compared with values obtained by inorganic precipitation experiments. This suggests a strong effect of cell physiology on biogenic calcite precipitation in coccolithophores. We present a conceptual model, based on the transmembrane transport of Sr and Ca, which explains the offset.

Description: Eutrophication can profoundly change rocky shore communities. These changes often cause the replacement of perennial, canopy-forming algae such as Fucus spp. with annual, bloom-forming algae such as Enteromorpha spp. Grazing, however, can counteract eutrophication by eliminating the annual algae�s susceptible recruits. We examine these generalizations across large scales. We use replicated ��bioassay�� experiments to compare the effects of eutrophication and grazing across four paired control versus eutrophied sites in the Northwest Atlantic and four eutrophied sites in the Baltic Sea in spring and summer. At each site, annual algal recruitment and grazing pressure were estimated using tiles seeded with Enteromorpha intestinalis propagules. Tiles were exposed for 3 weeks with grazers excluded or allowed access. Productivity of E. intestinalis recruits was strongly related to eutrophication (10-fold increase) and grazing (80% decrease) and was weakly related to season. While the absolute grazing rate increased in a linear fashion with algal productivity, the relative grazing rate remained surprisingly constant (;80%). Comparative field surveys showed that perennial algae decreased by 30-60%, while annual algae, filter feeders, and grazers increased across a gradient of eutrophication. As eutrophication increased from control to eutrophied to point source sites, rocky shore communities became increasingly dominated by single species of annual algae or filter feeders, and community diversity declined consistently by 24-46%. We conclude that grazers are important controllers of algal blooms but that, ultimately, they cannot override the effects of increasing eutrophication on rocky shore community structure and biodiversity.

Description: We investigated the effect of seasonal environmental changes on the rate and distribution of anaerobic oxidation of methane (AOM) in Eckernforde Bay sediments (German Baltic Sea) and identified organisms that are likely to be involved in the process. Surface sediments were sampled during September and March. Field rates of AOM and sulfate reduction (SR) were measured with radiotracer methods. Additional parameters were determined that potentially influence AOM, i.e., temperature, salinity, methane, sulfate, and chlorophyll a. Methanogenesis as well as potential rates of AOM and aerobic oxidation of methane were measured in vitro. AOM changed seasonally within the upper 20 cm of the sediment, with rates being between 1 and 14 nmol cm(-3) d(-1). Its distribution is suggested to be controlled by oxygen and sulfate penetration, temperature, as well as methane supply, leading to a shallow AOM zone during the warm productive season and to a slightly deeper AOM zone during the cold winter season. Rising methane bubbles apparently fed AOM above the sulfate-methane transition. Methanosarcinales-related anaerobic methanotrophs (ANME-2), identified with fluorescence in situ hybridization, is suggested to mediate AOM in Eckerntorde Bay. These archaea are known also from other marine methane-rich locations. However, they were not directly associated with sulfate-reducing bacteria. AOM is possibly mediated solely by these archaea that show a mesophilic physiology according to the seasonal temperature changes in Eckernforde Bay.

Description: We studied the direct effects of CO2 Engel et al. in the seawater was modified by an aeration system. The triplicate mesocosm treatments represented low (190 parts per million by volume (ppmV) CO2 organisms in a mesocosm experiment. In nine outdoor enclosures (11 m3 present (410 ppmV CO2 and high (710 ppmV CO2 pCO2 and related changes in seawater carbonate chemistry on marine planktonic each), the partial pressure of CO2 (pCO2 conditions. After initial fertilization with nitrate and phosphate a bloom dominated by the coccolithophorid Emiliania huxleyi occurred simultaneously in all of the nine mesocosms; it was monitored over a 19-day period. The three CO2 treatments assimilated nitrate and phosphate similarly. The concentration of particulate constituents was highly variable among the replicate mesocosms, disguising direct CO2 within each treatment, however, indicated that the net specific growth rate of E. huxleyi, the rate of calcification per cell, and the elemental stoichiometry of uptake and production processes were sensitive to changes in pCO2 This broad influence of CO2 -related effects. Normalization of production rates . on the E. huxleyi bloom suggests that changes in CO2 physiology with likely effects on the marine biogeochemistry.

Description: At Hook Ridge hydrothermal vent, a new species of Sclerolinum (Monilifera, Siboglinidae) was found at a water depth of 1,045 m. On the basis of investigations of multicores and gravity cores, the species habitat is characterized. Sclerolinum does not occur in sediments that are most strongly influenced by hydrothermal fluids, probably because of high temperature (up to 49°C) and precipitation of siliceous crusts. About 800 individuals m-2 occur in sediments that are only weakly exposed to hydrothermal flow and have the following characteristics: 20°C (15 cm sediment depth) to 21.5°C (bottom water), 18-40 cm yr-1 advection rates, pH 5.5, 〈25 µmol L-1 methane, 〈170 µmol L-1 sulfide, and 〈0.0054 mol m-2 yr-1 sulfide flux. Comparison with geochemical data from other reducing sediments indicates that the two groups of Siboglinidae, Monilifera and Frenulata, occur in sediments with low sulfide concentration and flux. In contrast, sulfurbased chemosynthetic organisms that typically occur at hydrothermal vents and cold seeps (e.g., Vestimentifera, vesicomyid clams, and bacterial mats) occur in sediments with higher sulfide availability; threshold values are around 500 µmol L-1 sulfide and 0.1 mol m-2 yr-1 sulfide fluxes. We did not find typical hydrothermal vent species at Hook Ridge hydrothermal vent, which might be explained by the unfavorable physicochemical habitat: At sites inhabited by Sclerolinum, sulfide availability appears to be too low, whereas at sites with higher sulfide availability, the temperatures might be too high, siliceous crust precipitation could preclude their occurrence, or both.