Chlorophyll-a, primary production rates, carbon concentrations and cell counts of coccolithophores, diatoms and microzooplankton measured in water samples from the North Atlantic during the M87/1 cruise in April 2012

The Deep Convection cruise repeatedly sampled two locations in the North Atlantic, sited in the Iceland and Norwegian Basins, onboard the RV Meteor (19 March - 2 May 2012). Samples were collected from multiple casts of a conductivity-temperature-depth (CTD) - Niskin rosette at each station. Water samples for primary production rates, community structure, chlorophyll a [Chl a], calcite [PIC], particulate organic carbon [POC] and biogenic silicic acid [BSi] were collected from predawn casts from six light depths (55%, 20%, 14%, 7%, 5% and 1% of incident PAR). Additional samples for community structure and ancillary parameters were collected from a second cast. Carbon fixation rates were determined using the 13C stable isotope method. Water samples for diatom and micro zooplankton counts, collected from the predawn casts, were preserved with acidic Lugol's solution (2% final solution) and counted using an inverted light microscope. Water samples for coccolithophore counts were collected onto cellulose nitrate filters and counted using polarising light microscopy. Water samples for Chl a analysis were filtered onto MF300 and polycarbonate filters and extracted in 90% acetone. PIC and BSi samples were filtered onto polycarbonate filters and analysed using an inductively coupled plasma emission optical spectrometer and a SEAL QuAAtro autoanalyser respectively.

Terrigenous input and marine productivity for the Pacific sector of the Southern Ocean based on lipid biomarkers

In this study, we present a new multiproxy data set of terrigenous input, marine productivity and sea surface temperature (SST) from 52 surface sediment samples collected along E-W transects in the Pacific sector of the Southern Ocean. Allochtonous terrigenous input was characterized by the distribution of plant wax n-alkanes and soil-derived branched glycerol dialkyl glycerol tetraethers (brGDGTs). 230Th-normalized burial rates of both compound groups were highest close to the potential sources in Australia and New Zealand and are strongly related to lithogenic contents, indicating common sources and transport. Detection of both long-chain n-alkanes and brGDGTs at the most remote sites in the open ocean strongly suggests a primarily eolian transport mechanism to at least 110°W, i.e. by prevailing westerly winds. Two independent organic SST proxies were used, the UK'37 based on long-chain alkenones, and the TEX86 based on isoprenoid GDGTs. Both, UK'37 and TEX86 indices show robust relationships with temperature over a temperature range between 0.5 and 20°C, likely implying different seasonal and regional imprints on the temperature signal. While alkenone-based temperature estimates reliably reflect modern SST even at the low temperature end, large temperature residuals are observed for the polar ocean using the TEX86 index. 230Th-normalized burial rates of alkenones are highest close to the Subtropical Front and are positively related to lithogenic fluxes throughout the study area. In contrast, highest isoGDGT burial south of the Antarctic Polar Front is not related with dust flux but may be largely controlled by diatom blooms, and thus high opal fluxes during austral summer.

Global distributions of diazotrophs nitrogen fixation rates - Depth integrated values computed from a collection of source datasets - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

The MAREDAT atlas covers 11 types of plankton, ranging in size from bacteria to jellyfish. Together, these plankton groups determine the health and productivity of the global ocean and play a vital role in the global carbon cycle. Working within a uniform and consistent spatial and depth grid (map) of the global ocean, the researchers compiled thousands and tens of thousands of data points to identify regions of plankton abundance and scarcity as well as areas of data abundance and scarcity. At many of the grid points, the MAREDAT team accomplished the difficult conversion from abundance (numbers of organisms) to biomass (carbon mass of organisms). The MAREDAT atlas provides an unprecedented global data set for ecological and biochemical analysis and modeling as well as a clear mandate for compiling additional existing data and for focusing future data gathering efforts on key groups in key areas of the ocean. The present data set presents depth integrated values of diazotrophs nitrogen fixation rates, computed from a collection of source data sets.

(Table 1) Major element composition of core 5022-2CP, Laguna Potrok Aike, southern Patagonia

Total organic carbon, total inorganic carbon, biogenic silica content and total organic carbon/total nitrogen ratios of the Laguna Potrok Aike lacustrine sediment record are used to reconstruct the environmental history of south-east Patagonia during the past 51 ka in high resolution. High lake level conditions are assumed to have prevailed during the Last Glacial, as sediments are carbonate-free. Increased runoff linked to permafrost and reduced evaporation due to colder temperatures and reduced influence of Southern Hemispheric Westerlies (SHW) may have caused these high lake levels with lake productivity being low and organic matter mainly of algal or cyanobacterial origin. Aquatic moss growth and diatom blooms occurred synchronously with southern hemispheric glacial warming events such as the Antarctic A-events, the postglacial warming following the LGM and the Younger Dryas chronozone. During these times, a combination of warmer climatic conditions with related thawing permafrost could have increased the allochthonous input of nutrients and in combination with warmer surface waters increased aquatic moss growth and diatom production. The SHW were not observed to affect southern Patagonia during the Last Glacial. The Holocene presents a completely different lacustrine system because (a) permafrost no longer inhibits infiltration nor emits meltwater pulses and (b) the positioning of the SHW over the investigated area gives rise to strong and dry winds. Under these conditions total organic carbon, total organic carbon/total nitrogen ratios and biogenic silica cease to be first order productivity indicators. On the one hand, the biogenic silica is influenced by dissolution of diatoms due to higher salinity and pH of the lake water under evaporative stress characterizing low lake levels. On the other hand, total organic carbon and total organic carbon/total nitrogen profiles are influenced by reworked macrophytes from freshly exposed lake level terraces during lowstands. Total inorganic carbon remains the most reliable proxy for climatic variations during the Holocene as high precipitation of carbonates can be linked to low lake levels and high autochthonous production. The onset of inorganic carbon precipitation has been associated with the southward shift of the SHW over the latitudes of Laguna Potrok Aike. The refined age-depth model of this record suggests that this shift occurred around 9.4 cal. ka BP.

Global distributions of diazotrophs abundance, biomass and nitrogen fixation rates - Gridded data product (NetCDF) - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

The MAREDAT atlas covers 11 types of plankton, ranging in size from bacteria to jellyfish. Together, these plankton groups determine the health and productivity of the global ocean and play a vital role in the global carbon cycle. Working within a uniform and consistent spatial and depth grid (map) of the global ocean, the researchers compiled thousands and tens of thousands of data points to identify regions of plankton abundance and scarcity as well as areas of data abundance and scarcity. At many of the grid points, the MAREDAT team accomplished the difficult conversion from abundance (numbers of organisms) to biomass (carbon mass of organisms). The MAREDAT atlas provides an unprecedented global data set for ecological and biochemical analysis and modeling as well as a clear mandate for compiling additional existing data and for focusing future data gathering efforts on key groups in key areas of the ocean. This is a gridded data product about diazotrophic organisms . There are 6 variables. Each variable is gridded on a dimension of 360 (longitude) * 180 (latitude) * 33 (depth) * 12 (month). The first group of 3 variables are: (1) number of biomass observations, (2) biomass, and (3) special nifH-gene-based biomass. The second group of 3 variables is same as the first group except that it only grids non-zero data. We have constructed a database on diazotrophic organisms in the global pelagic upper ocean by compiling more than 11,000 direct field measurements including 3 sub-databases: (1) nitrogen fixation rates, (2) cyanobacterial diazotroph abundances from cell counts and (3) cyanobacterial diazotroph abundances from qPCR assays targeting nifH genes. Biomass conversion factors are estimated based on cell sizes to convert abundance data to diazotrophic biomass. Data are assigned to 3 groups including Trichodesmium, unicellular diazotrophic cyanobacteria (group A, B and C when applicable) and heterocystous cyanobacteria (Richelia and Calothrix). Total nitrogen fixation rates and diazotrophic biomass are calculated by summing the values from all the groups. Some of nitrogen fixation rates are whole seawater measurements and are used as total nitrogen fixation rates. Both volumetric and depth-integrated values were reported. Depth-integrated values are also calculated for those vertical profiles with values at 3 or more depths.

Experiment: Response of Nodularia spumigena to pCO2

The filamentous and diazotrophic cyanobacterium Nodularia spumigena plays a major role in the productivity of the Baltic Sea as it forms extensive blooms regularly. Under phosphorus limiting conditions Nodularia spumigena has a high enzyme affinity for dissolved organic phosphorus (DOP) by production and release of alkaline phosphatase. Additionally, it is able to degrade proteinaceous compounds by expressing the extracellular enzyme leucine aminopeptidase. As atmospheric CO2 concentrations are increasing, we expect marine phytoplankton to experience changes in several environmental parameters including pH, temperature, and nutrient availability. The aim of this study was to investigate the combined effect of CO2-induced changes in seawater carbonate chemistry and of phosphate deficiency on the exudation of organic matter, and its subsequent recycling by extracellular enzymes in a Nodularia spumigena culture. Batch cultures of Nodularia spumigena were grown for 15 days aerated with three different pCO2 levels corresponding to values from glacial periods to future values projected for the year 2100. Extracellular enzyme activities as well as changes in organic and inorganic compound concentrations were monitored. CO2 treatment-related effects were identified for cyanobacterial growth, which in turn was influencing exudation and recycling of organic matter by extracellular enzymes. Biomass production was increased by 56.5% and 90.7% in the medium and high pCO2 treatment, respectively, compared to the low pCO2 treatment and simultaneously increasing exudation. During the growth phase significantly more mucinous substances accumulated in the high pCO2 treatment reaching 363 µg Gum Xanthan eq /l compared to 269 µg Gum Xanthan eq /l in the low pCO2 treatment. However, cell-specific rates did not change. After phosphate depletion, the acquisition of P from DOP by alkaline phosphatase was significantly enhanced. Alkaline phosphatase activities were increased by factor 1.64 and 2.25, respectively, in the medium and high compared to the low pCO2 treatment. In conclusion, our results suggest that Nodularia spumigena can grow faster under elevated pCO2 by enhancing the recycling of organic matter to acquire nutrients.

(Table 3) Interannual variability of total mass flux at the deep sediment trap site CBmeso between 1988 and 2006

This article will review major features of the 'giant' Cape Blanc filament off Mauritania with regard to the transport of chlorophyll and organic carbon from the shelf to the open ocean. Within the filament, chlorophyll is transported about 400 km offshore. Modelled particle distributions along a zonal transect at 21°N showed that particles with a sinking velocity of 5 m d**-1 are advected offshore by up to 600 km in subsurface particle clouds generally located between 400 m and 800 m water depth, forming an Intermediate Nepheloid Layer (INL). It corresponds to the depth of the oxygen minimum zone. Heavier particles with a sinking velocity of 30 m d**-1 are transported from the shelf within the Bottom Layer (BL) of more than 1000 m thickness, largely following the topography of the bottom slope. The particles advected within the BL contribute to the enhanced winter-spring mass fluxes collected at the open-ocean mesotrophic sediment trap site CB-13 (200 nm offshore), due to a long distance advection in deeper waters. The lateral contribution to the deep sediment trap in winter-spring is estimated to be 63% and 72% for organic carbon and total mass, respectively, whereas the lateral input for both components on an annual basis is estimated to be in the order of 15%. Biogenic opal increases almost fivefold from the upper to the lower mesotrophic CB-13 trap, also pointing to an additional source for biogenic silica from eutrophic coastal waters. Blooms obviously sink in smaller, probably mesoscale-sized patches with variable settling rates, depending on the type of aggregated particles and their ballast content. Generally, particle sinking rates are exceptionally high off NW Africa. Very high chlorophyll values and a large size of the Cape Blanc filament in 1998-1999 are also documented in enhanced total mass and organic carbon fluxes. An increasing trend in satellite chlorophyll concentrations and the size of the Cape Blanc filament between 1997 and 2008 as observed for other coastal upwelling areas is not documented.

(Table T2) Biogenic opal in Palmer Deep sediments at ODP Site 178-1098

High-resolution records of sedimentary proxies provide insights into fine-scale geochemical responses to climatic forcing. Gamma-ray attenuation (GRA) bulk-density data and magnetic stratigraphy records from Palmer Deep, Site 1098, show variability close to the same scale as ice cores, making this site ideal for high-resolution geochemical investigations. In conjunction with shipboard geophysical measurements, silica records allow high-resolution evaluation of the frequencies and amplitudes of biogenic variability. This provides investigators additional data sets to evaluate the global extent of climatic events that are presently defined by regional oceanic data sets (e.g., Younger Dryas in the North Atlantic) and to evaluate the potential mechanisms that link biological productivity and climate in the Southern Ocean. In addition, because of the observed links between diatom blooms and export productivity (Michaels and Silver, 1988, doi:10.1016/0198-0149(88)90126-4), biogenic silica may be an indicator of the efficiency of the biological pump (removal of organic carbon from the euphotic zone and burial within the sediments). Because the net removal of CO2 (on short time scales up to millennial, the balance between upwelled CO2, carbon fixation, and the removal of organic carbon from the surface ocean) can determine the atmospheric concentration; proxies that allow us to quantify export production yield insights into carbon cycle responses. In today's ocean, diatoms are integrally linked with new production (production based on the use of nitrate and molecular nitrogen rather than ammonium, which is generated by the microbial degradation of organic carbon) (Dugdale and Goering, 1967). Thus, as with nutrient utilization proxies, biogenic silica may be a good indicator of export production. The difficulties lie in translating the biogenic opal burial records to export production. Numerous factors control the preservation of sedimentary biogenic silica, including depth of the water column, water temperature, trace element chemistry, grazing pressure, bloom structure, and species composition of the diatom assemblage (Nelson et al., 1995, doi:10.1029/95GB01070). In addition, several recent investigations have noted additional complications. Iron limitation increases the uptake of Si relative to carbon (Hutchins et al., 1998, ; Takeda, 1998, doi:10.1038/31674). In the Southern Ocean, iron limitation could produce more robust, and thus better preserved, diatoms; thus, the burial record may be a record of iron limitation rather than of the export of organic carbon (Boyle, 1998). In addition, laboratory experiments show that bacteria accelerate the dissolution of biogenic silica (Bidle and Azam, 1999, doi:10.1038/17351). Both the species composition and temperature seem to influence the amount of dissolution. Evidence of recycling of silicic acid within the photic zone (Brzezinski et al., 1997) suggests that the silica pump (removal from the euphotic zone of silica relative to nitrogen and phosphorus) may work with variable efficiency. This becomes an issue when trying to reconstruct the removal of organic carbon from sedimentary biogenic silica records. In fact, there is a wide range in the Si:Corganic molar ratio in the Southern Ocean (0.18-0.81) (Nelson et al., 1995; Ragueneau et al., 2000, doi:10.1016/S0921-8181(00)00052-7). Thus, the presence (or absence) of biogenic silica alone may tell us little about the export productivity, complicating the interpretation of age-related trends. One recent assessment has added some hope to links between productivity and opal burial in the Southern Ocean (Pondaven et al., 2000). Quantitative comparison of different productivity proxies will greatly aid in this evaluation.

Warmer, deeper and greener mixed layers in the north Atlantic subpolar gyre over the last 50 years.

Shifts in global climate resonate in plankton dynamics, biogeochemical cycles, and marine food webs. We studied these linkages in the North Atlantic subpolar gyre (NASG), which hosts extensive phytoplankton blooms. We show that phytoplankton abundance increased since the 1960s in parallel to a deepening of the mixed layer and a strengthening of winds and heat losses from the ocean, as driven by the low frequency of the North Atlantic Oscillation (NAO). In parallel to these bottom-up processes, the top-down control of phytoplankton by copepods decreased over the same time period in the western NASG, following sea surface temperature changes typical of the Atlantic Multi-decadal Oscillation (AMO). While previous studies have hypothesized that climate-driven warming would facilitate seasonal stratification of surface waters and long-term phytoplankton increase in subpolar regions, here we show that deeper mixed layers in the NASG can be warmer and host a higher phytoplankton biomass. These results emphasize that different modes of climate variability regulate bottom-up (NAO control) and top-down (AMO control) forcing on phytoplankton at decadal timescales. As a consequence, different relationships between phytoplankton, zooplankton, and their physical environment appear subject to the disparate temporal scale of the observations (seasonal, interannual, or decadal). The prediction of phytoplankton response to climate change should be built upon what is learnt from observations at the longest timescales.

Warmer, deeper and greener mixed layers in the north Atlantic subpolar gyre over the last 50 years.

Shifts in global climate resonate in plankton dynamics, biogeochemical cycles, and marine food webs. We studied these linkages in the North Atlantic subpolar gyre (NASG), which hosts extensive phytoplankton blooms. We show that phytoplankton abundance increased since the 1960s in parallel to a deepening of the mixed layer and a strengthening of winds and heat losses from the ocean, as driven by the low frequency of the North Atlantic Oscillation (NAO). In parallel to these bottom-up processes, the top-down control of phytoplankton by copepods decreased over the same time period in the western NASG, following sea surface temperature changes typical of the Atlantic Multi-decadal Oscillation (AMO). While previous studies have hypothesized that climate-driven warming would facilitate seasonal stratification of surface waters and long-term phytoplankton increase in subpolar regions, here we show that deeper mixed layers in the NASG can be warmer and host a higher phytoplankton biomass. These results emphasize that different modes of climate variability regulate bottom-up (NAO control) and top-down (AMO control) forcing on phytoplankton at decadal timescales. As a consequence, different relationships between phytoplankton, zooplankton, and their physical environment appear subject to the disparate temporal scale of the observations (seasonal, interannual, or decadal). The prediction of phytoplankton response to climate change should be built upon what is learnt from observations at the longest timescales.

Effects of a harmful algal bloom on the community ecology, movements and spatial distributions of fishes in a microtidal estuary

Harmful algal blooms can adversely affect fish communities, though their impacts are highly context-dependent and typically differ between fish species. Various approaches, comprising univariate and multivariate analyses and multimetric Fish Community Indices (FCI), were employed to characterise the perceived impacts of a Karlodinium veneficum bloom on the fish communities and ecological condition of the Swan Canning Estuary, Western Australia. The combined evidence suggests that a large proportion of the more mobile fish species in the offshore waters of the bloom-affected area relocated to other regions during the bloom. This was indicated by marked declines in mean species richness, catch rates and FCI scores in the bloom region but concomitant increases in these characteristics in more distal regions, and by pronounced and atypical shifts in the pattern of inter-regional similarities in fish community composition during the bloom. The lack of any significant changes among the nearshore fish communities revealed that bloom impacts were less severe there than in deeper, offshore waters. Nearshore habitats, which generally are in better ecological condition than adjacent offshore waters in this system, may provide refuges for fish during algal blooms and other perturbations, mirroring similar observations of fish avoidance responses to such stressors in estuaries worldwide.

Effects of a harmful algal bloom on the community ecology, movements and spatial distributions of fishes in a microtidal estuary

Harmful algal blooms can adversely affect fish communities, though their impacts are highly context-dependent and typically differ between fish species. Various approaches, comprising univariate and multivariate analyses and multimetric Fish Community Indices (FCI), were employed to characterise the perceived impacts of a Karlodinium veneficum bloom on the fish communities and ecological condition of the Swan Canning Estuary, Western Australia. The combined evidence suggests that a large proportion of the more mobile fish species in the offshore waters of the bloom-affected area relocated to other regions during the bloom. This was indicated by marked declines in mean species richness, catch rates and FCI scores in the bloom region but concomitant increases in these characteristics in more distal regions, and by pronounced and atypical shifts in the pattern of inter-regional similarities in fish community composition during the bloom. The lack of any significant changes among the nearshore fish communities revealed that bloom impacts were less severe there than in deeper, offshore waters. Nearshore habitats, which generally are in better ecological condition than adjacent offshore waters in this system, may provide refuges for fish during algal blooms and other perturbations, mirroring similar observations of fish avoidance responses to such stressors in estuaries worldwide.