Gut content of holothurians from the NE Atlantic

This paper describes inter-specific differences in the distribution of sediment in the gut compartments and in the enzyme and bacterial profiles along the gut of abyssal holothurian species - Oneirophanta mutabilis, Psychropotes longicauda and Pseudostichopus villosus sampled from a eutrophic site in the NE Atlantic at different times of the year. Proportions of sediments, relative to total gut contents, in the pharynx, oesophagus, anterior and posterior intestine differed significantly in all the inter-species comparisons, but not between inter-seasonal comparisons. Significant differences were also found between the relative proportions of sediments in both the rectum and cloaca of Psychropotes longicauda and Oneirophanta mutabilis. Nineteen enzymes were identified in either gut-tissue or gut-content samples of the holothurians studied. Concentrations of the enzymes in gut tissues and their contents were highly correlated. Greater concentrations of the enzymes were found in the gut tissues suggesting that they are the main source of the enzymes. The suites of enzymes recorded were broadly similar in each of the species sampled collected regardless of the time of the year, and they were similar to those described previously for shallow-water holothurians. Significant inter-specific differences in the gut tissue concentrations of some of the glycosidases suggest dietary differences. For example, Psychropotes longicauda and Pseudostichopus villosus contain higher levels of chitobiase than Oneirophanta mutabilis. There were no seasonal changes in bacterial activity profiles along the guts of O. mutabilis and Pseudostichopus villosus. In both these species bacterial activity and abundance declined between the pharynx/oesophagus and anterior intestine, but then increased along the gut and became greatest in the rectum/cloaca. Although the data sets were more limited for Psychropotes longicauda, bacterial activity increased from the anterior to the posterior intestine but then declined slightly to the rectum/cloaca. These changes in bacterial activity and densities probably reflect changes in the microbial environment along the guts of abyssal holothurians. Such changes suggest that there is potential for microbial breakdown of a broader range of substrates than could be otherwise be achieved by the holothurian itself. However, the present study found no evidence for sedimentary (microbial) sources of hydrolytic enzymes.

Biological and environmental time series obtained from Point B, Bay of Villefranche

Ecological succession provides a widely accepted description of seasonal changes in phytoplankton and mesozooplankton assemblages in the natural environment, but concurrent changes in smaller (i.e. microbes) and larger (i.e. macroplankton) organisms are not included in the model because plankton ranging from bacteria to jellies are seldom sampled and analyzed simultaneously. Here we studied, for the first time in the aquatic literature, the succession of marine plankton in the whole-plankton assemblage that spanned 5 orders of magnitude in size from microbes to macroplankton predators (not including fish or fish larvae, for which no consistent data were available). Samples were collected in the northwestern Mediterranean Sea (Bay of Villefranche) weekly during 10 months. Simultaneously collected samples were analyzed by flow cytometry, inverse microscopy, FlowCam, and ZooScan. The whole-plankton assemblage underwent sharp reorganizations that corresponded to bottom-up events of vertical mixing in the water-column, and its development was top-down controlled by large gelatinous filter feeders and predators. Based on the results provided by our novel whole-plankton assemblage approach, we propose a new comprehensive conceptual model of the annual plankton succession (i.e. whole plankton model) characterized by both stepwise stacking of four broad trophic communities from early spring through summer, which is a new concept, and progressive replacement of ecological plankton categories within the different trophic communities, as recognised traditionally.

Particle size of Saharan dust in the Atlantic Ocean, from October 2012 to November 2013

Mineral dust has a large impact on regional and global climate, depending on its particle size. Especially in the Atlantic Ocean downwind of the Sahara, the largest dust source on earth, the effects can be substantial but are poorly understood. This study focuses on seasonal and spatial variations in particle size of Saharan dust deposition across the Atlantic Ocean, using an array of submarine sediment traps moored along a transect at 12° N. We show that the particle size decreases downwind with increased distance from the Saharan source, due to higher gravitational settling velocities of coarse particles in the atmosphere. Modal grain sizes vary between 4 and 33 µm throughout the different seasons and at five locations along the transect. This is much coarser than previously suggested and incorporated into climate models. In addition, seasonal changes are prominent, with coarser dust in summer, and finer dust in winter and spring. Such seasonal changes are caused by transport at higher altitudes and at greater wind velocities during summer than in winter. Also the latitudinal migration of the dust cloud, associated with the Intertropical Convergence Zone, causes seasonal differences in deposition as the summer dust cloud is located more to the north, and more directly above the sampled transect. Furthermore, increased precipitation and more frequent dust storms in summer coincide with coarser dust deposition. Our findings contribute to understanding Saharan dust transport and deposition relevant for the interpretation of sedimentary records for climate reconstructions, as well as for global and regional models for improved prediction of future climate.

Effects of in situ CO2 enrichment on structural characteristics, photosynthesis, and growth of the Mediterranean seagrass Posidonia oceanica

Seagrass is expected to benefit from increased carbon availability under future ocean acidification. This hypothesis has been little tested by in situ manipulation. To test for ocean acidification effects on seagrass meadows under controlled CO2/pH conditions, we used a Free Ocean Carbon Dioxide Enrichment (FOCE) system which allows for the manipulation of pH as continuous offset from ambient. It was deployed in a Posidonia oceanica meadow at 11 m depth in the Northwestern Mediterranean Sea. It consisted of two benthic enclosures, an experimental and a control unit both 1.7 m**3, and an additional reference plot in the ambient environment (2 m**2) to account for structural artifacts. The meadow was monitored from April to November 2014. The pH of the experimental enclosure was lowered by 0.26 pH units for the second half of the 8-month study. The greatest magnitude of change in P. oceanica leaf biometrics, photosynthesis, and leaf growth accompanied seasonal changes recorded in the environment and values were similar between the two enclosures. Leaf thickness may change in response to lower pH but this requires further testing. Results are congruent with other short-term and natural studies that have investigated the response of P. oceanica over a wide range of pH. They suggest any benefit from ocean acidification, over the next century (at a pH of 7.7 on the total scale), on Posidonia physiology and growth may be minimal and difficult to detect without increased replication or longer experimental duration. The limited stimulation, which did not surpass any enclosure or seasonal effect, casts doubts on speculations that elevated CO2 would confer resistance to thermal stress and increase the buffering capacity of meadows.

Synergistic effects of ocean acidification and warming on overwintering pteropods in the Arctic

Ocean acidification and warming will be most pronounced in the Arctic Ocean. Aragonite shell-bearing pteropods in the Arctic are expected to be among the first species to suffer from ocean acidification. Carbonate undersaturation in the Arctic will first occur in winter and because this period is also characterized by low food availability, the overwintering stages of polar pteropods may develop into a bottleneck in their life cycle. The impacts of ocean acidification and warming on growth, shell degradation (dissolution), and mortality of two thecosome pteropods, the polar Limacina helicina and the boreal L. retroversa, were studied for the first time during the Arctic winter in the Kongsfjord (Svalbard). The abundance of L. helicina and L. retroversa varied from 23.5 to 120 ind /m2 and 12 to 38 ind /m2, and the mean shell size ranged from 920 to 981 µm and 810 to 823 µm, respectively. Seawater was aragonite-undersaturated at the overwintering depths of pteropods on two out of ten days of our observations. A 7-day experiment [temperature levels: 2 and 7 °C, pCO2 levels: 350, 650 (only for L. helicina) and 880 ?atm] revealed a significant pCO2 effect on shell degradation in both species, and synergistic effects between temperature and pCO2 for L. helicina. A comparison of live and dead specimens kept under the same experimental conditions indicated that both species were capable of actively reducing the impacts of acidification on shell dissolution. A higher vulnerability to increasing pCO2 and temperature during the winter season is indicated compared with a similar study from fall 2009. Considering the species winter phenology and the seasonal changes in carbonate chemistry in Arctic waters, negative climate change effects on Arctic thecosomes are likely to show up first during winter, possibly well before ocean acidification effects become detectable during the summer season.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.

Concentrations and d13C compositions of n-alkanes, n-alcohols, n-alkanoic acids in a 34-month time series of suspended sediments from the outflow of the Congo River

The concentrations, distributions, and stable carbon isotopes (d13C) of plant waxes carried by fluvial suspended sediments contain valuable information about terrestrial ecosystem characteristics. To properly interpret past changes recorded in sedimentary archives it is crucial to understand the sources and variability of exported plant waxes in modern systems on seasonal to inter-annual timescales. To determine such variability, we present concentrations and d13C compositions of three compound classes (n-alkanes, n-alcohols, n-alkanoic acids) in a 34-month time series of suspended sediments from the outflow of the Congo River. We show that exported plant-dominated n-alkanes (C25-C35) represent a mixture of C3 and C4 end members, each with distinct molecular distributions, as evidenced by an 8.1 ± 0.7 per mil (±1Sigma standard deviation) spread in d13C values across chain-lengths, and weak correlations between individual homologue concentrations (r = 0.52-0.94). In contrast, plant-dominated n-alcohols (C26-C36) and n-alkanoic acids (C26-C36) exhibit stronger positive correlations (r = 0.70-0.99) between homologue concentrations and depleted d13C values (individual homologues average <= -31.3 per mil and -30.8 per mil, respectively), with lower d13C variability across chain-lengths (2.6 ± 0.6 per mil and 2.0 ± 1.1 per mil, respectively). All individual plant-wax lipids show little temporal d13C variability throughout the time-series (1 Sigma <= 0.9 per mil), indicating that their stable carbon isotopes are not a sensitive tracer for temporal changes in plant-wax source in the Congo basin on seasonal to inter-annual timescales. Carbon-normalized concentrations and relative abundances of n-alcohols (19-58% of total plant-wax lipids) and n-alkanoic acids (26-76%) respond rapidly to seasonal changes in runoff, indicating that they are mostly derived from a recently entrained local source. In contrast, a lack of correlation with discharge and low, stable relative abundances (5-16%) indicate that n-alkanes better represent a catchment-integrated signal with minimal response to discharge seasonality. Comparison to published data on other large watersheds indicates that this phenomenon is not limited to the Congo River, and that analysis of multiple plant-wax lipid classes and chain lengths can be used to better resolve local vs. distal ecosystem structure in river catchments.

Coastal plankton communities under elevated CO2: community barcoding results, link to data files in FASTA format

The acidification of the oceans could potentially alter marine plankton communities with consequences for ecosystem functioning. While several studies have investigated effects of ocean acidifications on communities using traditional methods, few have used genetic analyses. Here, we use community barcoding to assess the impact of ocean acidification on the composition of a coastal plankton community in a large scale, in situ, long-term mesocosm experiment. High-throughput sequencing resulted in the identification of a wide range of planktonic taxa (Alveolata, Cryptophyta, Haptophyceae, Fungi, Metazoa, Hydrozoa, Rhizaria, Straminipila, Chlorophyta). Analyses based on predicted operational taxonomical units as well as taxonomical compositions revealed no differences between communities in high CO2 mesocosms (~760 µatm) and those exposed to present day CO2 conditions. Observed shifts in the planktonic community composition were mainly related to seasonal changes in temperature and nutrients.

Stable isotope and Mg/Ca ratios of Globigerinoides ruber from core-top sediment samples

Seasonal changes in surface ocean temperature are increasingly recognized as an important parameter of the climate system. Here we assess the potential of analyzing single-specimen planktonic foraminifera as proxy for the seasonal temperature contrast (seasonality). Oxygen isotopes and Mg/Ca ratios were measured on single specimens of Globigerinoides ruber, extracted from surface sediment samples of the Mediterranean Sea and the adjacent Atlantic Ocean. Variability in d18O and Mg/Ca was then compared to established modern seasonal changes in temperature and salinity for both regions. The results show that (1) average d18O-derived temperatures correlate with modern annual average temperatures for most sites, (2) the range in d18O- and Mg/Ca-derived temperature estimates from single-specimen analysis resembles the range in seasonal temperature values at the sea surface (0-50 m) in the Mediterranean Sea and the Atlantic Ocean, and (3) there is no strong correlation between Mg/Ca- and d18O-derived temperatures from the same specimens in the current data set, indicating that other parameters (salinity, carbonate ion concentration, symbiont activity, ontogenesis, and natural variability) potentially affect these proxies.

Mean redox potential values in surface sediments along the transect D in the Cretan Sea (Table 1)

The seasonal, spatial and bathymetric changes in the distribution of chloroplastic pigments (Chl a, phaeopigments and CPE), TOC, TON, ATP, bottom water nutrient content and the main biochemical classes of organic compounds (lipids, proteins and carbohydrates) were recorded from May 1994 to September 1995 over the continental margin of northern Crete. The concentration of chloroplastic pigment equivalents (CPE) was always low, dropping dramatically along the shelf-slope gradient. Microbial activity (ATP) also dropped sharply beyond the continental shelf following a distribution pattern similar to TOC and TON. Lipid, protein and carbohydrate concentrations, as well as biopolymeric carbon were comparable to those reported for other more productive areas, however, the quality of the organic matter itself was rather poor. Thus, carbohydrates, the dominant biochemical class, were characterised by being highly (80-99%) refractory, as soluble carbohydrates represented (on annual average) only 6% of the total carbohydrate pool. Protein and lipid concentrations strongly decreased with depth, indicating depletion of trophic resources in the bathyal zone. Proteins appeared to be the more degradable compounds and indeed the protein to carbohydrate ratios were found to decrease strongly in the deeper stations. Organic matter content and quality decreased both with increasing distance from the coast and within the sediment. All sedimentary organic compounds were found to vary between sampling periods, with the changes being more pronounced over the continental shelf. The different temporal patterns of the various components suggest a different composition and/or origin of the OM inputs during the different sampling periods. The amount of material reaching the sediments below 540 m is extremely low, suggesting that most of the organic material is decomposed and/or utilised before reaching the sea floor. In conclusion, the continental shelf and bathyal sediments of the Cretan Sea can be considered, from a trophic point of view, as two different subsystems.

Meiofauna of surface sediments in the Cretan Sea

Quantitative information on metazoan meiofaunal abundance and biomass was obtained from three continental shelf (at 40, 100 and 200 m depth) and four deep-sea stations (at 540, 700, 940 and 1540 m depth) in the Cretan Sea (South Aegean Sea, NE Mediterranean). Samples were collected on a seasonal basis (from August 1994 to September 1995) with the use of a multiple corer. Meiofaunal abundance and biomass on the continental shelf of the Cretan Sea were high, in contrast to the extremely low values reported for the bathyal sediments that showed values comparable to those reported for abyssal and hadal environments. In order to explain the spatial and seasonal changes in metazoan meiofauna these data were compared with: (1) the concentrations of 'food indicators' (such as proteins, lipids, soluble carbohydrates and CPE) (2) the bacterial biomass (3) the flux of labile organic compounds to the sea floor at a fixed station (D7, 1540 m depth). Highly significant relationships between meiofaunal parameters and CPE, protein and lipid concentrations and bacterial biomass were found. Most of the indicators of food quality and quantity (such as CPE, proteins and carbohydrates) showed a clear seasonality with highest values in February and lowest in September. Such changes were more evident on the continental shelf rather than at deeper depths. On the continental shelf, significant seasonal changes in meiofaunal density were related to changes in the input of labile organic carbon whereas meiofaunal assemblages on the deep-sea stations showed time-lagged changes in response to the food input recorded in February 95. At all deep-sea stations meiofaunal density increased with a time lag of 2 months. Indications for a time-lagged meiofaunal response to the food inputs were also provided by the increase in nauplii densities during May 95 and the increase in individual biomass of nematodes, copepods and polychaetes between February and May 1995. The lack of strong seasonal changes in deep sea meiofaunal density suggests that the supply of organic matter below 500 m is not strong enough to support a significant meiofaunal development. Below 700 m depth >92% of the total biomass in the sediment was represented by bacteria. The ratio of bacterial to meiofaunal biomass increased with increasing water depth indicating that bacteria are probably more effective than meiofauna in exploiting refractory organic compounds. These data lead us to hypothesise that the deep-sea sediments of the Cretan Sea are largely dependent upon a benthic microbial loop.

Nutrient availability affects the response of the calcifying chlorophyte Halimeda opuntia (L.) J.V. Lamouroux to low pH

Atmospheric carbon dioxide emissions cause a decrease in the pH and aragonite saturation state of surface ocean water. As a result, calcifying organisms are expected to suffer under future ocean conditions, but their physiological responses may depend on their nutrient status. Because many coral reefs experience high inorganic nutrient loads or seasonal changes in nutrient availability, reef organisms in localized areas will have to cope with elevated carbon dioxide and changes in inorganic nutrients. Halimeda opuntia is a dominant calcifying primary producer on coral reefs that contributes to coral reef accretion. Therefore, we investigated the carbon and nutrient balance of H. opuntia exposed to elevated carbon dioxide and inorganic nutrients. We measured tissue nitrogen, phosphorus and carbon content as well as the activity of enzymes involved in inorganic carbon uptake and nitrogen assimilation (external carbonic anhydrase and nitrate reductase, respectively). Inorganic carbon content was lower in algae exposed to high CO2, but calcification rates were not significantly affected by CO2 or inorganic nutrients. Organic carbon was positively correlated to external carbonic anhydrase activity, while inorganic carbon showed the opposite correlation. Carbon dioxide had a significant effect on tissue nitrogen and organic carbon content, while inorganic nutrients affected tissue phosphorus and N:P ratios. Nitrate reductase activity was highest in algae grown under elevated CO2 and inorganic nutrient conditions and lowest when phosphate was limiting. In general, we found that enzymatic responses were strongly influenced by nutrient availability, indicating its important role in dictating the local responses of the calcifying primary producer H. opuntia to ocean acidification.

Geochemical and stable isotope composition of Middle Campanian marl-limestone rhythmites of the Lehrte West Syncline near Hannover (Lower Saxony, northern Germany)

A cyclic marl-limestone succession of Middle-Late Campanian age has been investigated with respect to a Milankovitch-controlled origin of geochemical data. In general, the major element geochemistry of the marl-limestone rhythmites can be explained by a simple two-component mixing model with the end-members calcium carbonate and 'average shale'-like material. Carbonate content varies from 55 to 90%. Non-carbonate components are clay minerals (illite, smectite) and biogenic silica from sponge spicules, as well as authigenically formed zeolites (strontian heulandite) and quartz. The redox potential suggests oxidizing conditions throughout the section. Trace element and stable isotopic data as well as SEM investigations show that the carbonate mud is mostly composed of low-magnesium calcitic tests of planktic coccolithophorids and calcareous dinoflagellate cysts (calcispheres). Diagenetic overprint results in a decrease of 2% d18O and an increase in Mn of up to 250 ppm. However, the sediment seems to preserve most of its high Sr content compared to the primary low-magnesium calcite of co-occurring belemnite rostra. The periodicity of geochemical cycles is dominated by 413 ka and weak signals between 51 and 22.5 ka, attributable to orbital forcing. Accumulation rates within these cycles vary between 40 and 50 m/Ma. The resulting cyclic sedimentary sequence is the product of (a) changes in primary production of low-magnesium calcitic biogenic material in surface waters within the long eccentricity and the precession, demonstrated by the CaCO3 content and the Mg/Al, Mn/Al and Sr/Al ratios, and (b) fluctuations in climate and continental weathering, which changed the quality of supplied clay minerals (the illite/smectite ratio), demonstrated by the K/Al ratio. High carbonate productivity correlates with smectite-favouring weathering (semi-arid conditions, conspicuously dry and moist seasonal changes in warmer climates). Ti as the proxy indicator for the detrital terrigenous influx, as well as Rb, Si, Zr and Na, shows only low frequency signals, indicating nearly constant rates of supply throughout the more or less pure pelagic carbonate deposition of the long-lasting third-order Middle-Upper Campanian sedimentary cycle.