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Highlights •A solid knowledgebase and governance process is required to develop policies for sustainable growth of offshore aquaculture in the North Sea. •Environmental goals largely define maximum sustainable production levels. For seaweed culture this is limited to the scale of several hundred km2. •Definition of realistic ambitions is a process rather than a fixed value and greatly benefits from a ‘learning by doing’ approach.

The cryptophyte Rhodomonas salina is widely used in aquaculture due to its high nutritional profile. This study aims to investigate the effect of salinity and pH on the growth, phycoerythrin concentrations, and concentrations of non-volatile umami taste active compounds of R. salina, using a design of experiment approach. Rhodomonas salina was cultivated in a flat-panel photobioreactor in turbidostat mode in a range of salinity (20–40 ‰) and pH (6.5–8.5). The strain was able to grow steadily under all conditions, but the optimal productivity of 1.17 g dry weight L−1 day−1 was observed in salinity 30 ‰ and pH 7.5. The phycoerythrin concentration was inversely related to productivity, presenting higher values in conditions that were not optimal for the growth of R. salina, 7% of dry weight at salinity 40 ‰, and pH 8.5. The identification of the umami taste of R. salina was based on the synergistic effect of umami compounds 5′-nucleotides (adenosine 5′-monophosphate, guanosine 5′-monophosphate, inosine 5′-monophosphate) and free amino acids (glutamic and aspartic acids), using the equivalent umami concentration (EUC). The results indicated that an increase in pH induces the accumulation of 5′-nucleotides, resulting in an EUC of 234 mg MSG g−1 at a salinity of 40 and pH 8.5. The EUC values that were observed in R. salina were higher compared to other aquatic animals, a fact that makes R. salina promising for further research and application in the food and feed sectors.

Morphology and physiology are two key aspects of the adaptation of kelp to varying environments. Some of these kelp responses to co-occurring highly hydrodynamic condition and high nutrient availability are well documented, but little is known about how these factors affect frond surface shape, particularly in the central frond. In this study, morphological and physiological traits of acclimatized Saccharina latissima (Phaeophyceae) (three size classes: 44.14 ± 1.15 cm, 29.60 ± 0.75 cm, and 16.07 ± 0.45 cm) were compared after 56 days under fully controlled conditions of waves or no waves, and high or low nutrient availability (i.e., LN-NW, LN-W, HN-NW, and HN-W treatments). Waves primarily increased frond biomass, elongation rate, and carbon to nitrogen ratio (C:N ratio) and induced both a greater variety in and rougher frond surface shapes. The fastest, second-fastest, and slowest growth rates were observed in the HN-W, LN-W, and LN-NW treatments, respectively. The highest C:N ratio was observed in the LN-W treatment. Together, these results seem to suggest that the thready and spring-like shapes found in the central frond (i.e., rougher frond surface) in wave-exposed conditions can at least partly compensate for low nutrient availability by enhancing nutrient and photon acquisition, particularly in low nutrient conditions. Additionally, large individuals showed significantly larger and heavier fronds compared with other size classes, and the meristematic sections of fronds had the most variance in frond surface shapes and highest C:N ratios compared with distal and mid-sections. Together, these results indicate that frond surface shapes in the newly formed central frond of S. latissima can be regarded both as possessing high morphological and physiological plasticity that enables kelp to cope with contrasting environments.

Seaweed phytosterols are associated with potential health benefits, affording them and the seaweeds that produce them commercial interest. However, little is known about how their production is affected by the cultivation environment, limiting the efficiency with which these compounds can be exploited. Therefore, we performed a pilot study on the effect of nutrient availability and light stress on the sterol content of Saccharina latissima, a rapid growing brown alga of increasing interest in western mariculture. Individuals of S. latissima were subjected to a nutrient-replete and nutrient-depleted regime for 5 weeks, followed by the introduction of light-limited and light-saturated conditions in the sixth week; sampling occurred each week. No significant inter-treatment differences were found in the sterol content in week 1–5. However, significant intra-treatment differences were found in weeks 3–5 regardless of nutrient treatment, wherein the fucosterol, 24-methylenecholesterol, and squalene contents of both treatment groups were found to correlate inversely with photosynthetic performance. Factorial treatment of differential nutrient availability and light stress resulted in marked differences between the sterol content of all groups in week 6. Here, squalene and cycloartenol increased in concentration with increasing irradiance regardless of nutrient treatment. Concentrations of all other sterolic components increased with increasing irradiance and low nutrient conditions while decreasing or remaining unchanged with increasing irradiance and high nutrient conditions. Our data shows that within our cultivation conditions and time frame, the sterol content of S. latissima is unaffected by nutrient availability alone but changes with combined alterations in irradiance and nutrient availability. Graphical abstract

Uptake dynamics of dissolved inorganic phosphate (DIP) and dissolved inorganic nitrate (DIN) in young Palmaria palmata (n = 49), cultivated in a range of DIP concentrations (0.0–6.0 µmol · L−1) and nonlimiting DIN concentration (50 µmol · L−1) under fully controlled laboratory conditions, were quantified in a ‘pulse‐and‐chase’ approach over 5 weeks. Two different uptake rates were specified: (1) surge uptake (VS) after starvation and (2) maintenance uptake with filled nutrient pools (VMS−2 · d−1 and DIN of 15.6 ± 4.3 µmol · cm−2 · d−1 , as well as VM−2 · d−1 and DIN of 5.6 ± 2.1 µmol · cm−2 · d−1 were calculated. In addition, an absolute size of the internal storage capacity (ISC) for DIP of 22 µmol · cm2 and DIN of 222 µmol · cm2 was determined. A DIP‐to‐DIN uptake ratio of 1:10 under VMSP. palmata and allow for an optimization in cultivation.

Rhodomonas salina is a microalgal species, belonging to the cryptophytes, and is widely used as aquaculture feed because of its high nutritional profile and phycoerythrin content. This study investigated the effect of pH on the growth, biochemical composition, and taste of R. salina when cultivated on a semi-large scale under natural light conditions. Two tubular photobioreactors (200 L) were used for the cultivation of R. salina with sunlight as the only illumination source. Two different pH setpoints were applied, 7 and 8.5. Optimal temperature and nutrient conditions were applied, according to previous research findings. The results demonstrated that the productivity of R. salina was higher at pH 7, 0.06–0.14 gdry weight L−1 day−1, compared to pH 8.5, 0.03–0.12 gdry weight L−1 day−1. It was found that protein and total fatty acid concentrations were higher in the biomass that was produced at pH 8.5, 33.7% and 12.3% of dry weight, respectively, while at pH 7, the protein content was 31.9% and the total fatty acids 8.8% of dry weight. The phycoerythrin concentration, like protein, was higher at pH 8.5, 2.7% of dry weight, compared to pH 7, 1% of dry weight. The free amino acid and nucleotide profile of R. salina was affected by the pH, resulting in increased equivalent umami concentration at pH 7. For the sensory evaluation, an expert panel on algae flavors evaluated the effect of pH on the taste of R. salina, reporting that the biomass that was produced at pH 7 had more umami flavor than the biomass that was produced at pH 8.5, which was evaluated as more bitter.

Cultivation of Saccharina latissima, a brown macroalga, is fast developing in Europe and North America for the sustainable production of food and biorefinery materials and important ecosystem services. Prior studies have demonstrated large spatial variability in the yield and chemical composition of the cultivated S. latissima, even within a small coastal bay. Using a validated hydrodynamic-biogeochemical-kelpmodel, this study examined main drivers of the spatial variability in S. latissima growth dynamics in 40 hypothetical farms throughout a Dutch coastal bay, the Eastern Scheldt. Results indicate that temperatureplays a primary role in driving the spatial variability. For example, S. latissima yield in the deeper and better flushed western part is more than double that in the eastern part, mainly due to its 2–3°C warmer seawater in winter. It is also found that S. latissima benefits from co-cultivation with shellfish, since nutrients excreted by shellfish replenish its nitrogen reserve, which fuels a relatively high growth rate in the nitrogen-depleted late spring. The model assessment offers insight into optimal potential locations of S. latissima farms in the Eastern Scheldt. Applicability of our modelling approach to other coastal ecosystems and possible further improvements for assisting in seaweed farming practice are discussed.

Assessing the umami taste of seaweed on a chemical level can inform the use and selection of seaweed in European cuisine. Accordingly, we developed a method for the simultaneous extraction, separate clean-up and analysis of 21 free amino acids and 10 free nucleotides by reversed phase and mixed-mode HPLC respectively. Of multiple mouth emulating solvents, extracting in Milli-Q at 35 °C was found most suitable. This method showed good linearity (R2 > 0.9996), resolution (Rs ≥ 1.5) and picomole detection limits, and was successfully applied to determine the Equivalent Umami Concentration (EUC) and Taste Activity Values (TAV) of seven Dutch seaweed species. Phaeophyceae showedthe highest EUC, followed by Chlorophyceae and Rhodophyceae (≈ 9.5, 3.7 and 1.1 g/100 g respectively). Glutamic acid always exceeded the TAV, while other umami compounds were species specific. Our method can accurately predict umami intensity and therefore contributes towards species selection for the European palette.

Kelp cultivation receives increasing interest for its high-value products and ecological services, especially in Europe and North America. Before industrial kelp farming in marine ecosystems continue to scale up, evaluation of the site-wide production relative to ecological carrying capacity (CC) of the identified system is essential. For this purpose, a mechanistic kelp model was developed and applied for hypothetical numerical experiments of expanding the farming area in a Dutch coastal bay (the Eastern Scheldt), where cultivation of Saccharina latissima (sugar kelp) is emerging. The kelp model was implemented within a three-dimensional hydrodynamic–biogeochemical model to account for the environmental interactions. The model captured the seasonal growth dynamics of S. latissima, as well as its carbon and nitrogen contents measured at the Eastern Scheldt pilot sites. The model results suggest that expanding the kelp farming area to ∼1–30% of the bay (representing ∼3.4–75 kt harvest dry weight in the 350-km2 bay) had the potential to weaken the spring bloom, and thereby affected the coexisting shellfish culture in the bay. Competition between S. latissima and phytoplankton mostly occurred in late spring for nutrients (dissolved inorganic nitrogen). The ecological CC should be weighed according to these negative impacts. However, the production CC was not reached even when farming ∼30% of the Eastern Scheldt, i.e. harvesting totally 75 kt dry mass, given that the simulated overall S. latissima production kept increasing with the farming activity. Our modelling approach can be applied to other systems for S. latissima cultivation and assist in assessing CC and environmental impacts.

Eukaryotic phytoplankton are responsible for at least 20% of annual global carbon fixation. Their diversity and activity are shaped by interactions with prokaryotes as part of complex microbiomes. Although differences in their local species diversity have been estimated, we still have a limited understanding of environmental conditions responsible for compositional differences between local species communities on a large scale from pole to pole. Here, we show, based on pole-to-pole phytoplankton metatranscriptomes and microbial rDNA sequencing, that environmental differences between polar and non-polar upper oceans most strongly impact the large-scale spatial pattern of biodiversity and gene activity in algal microbiomes. The geographic differentiation of co-occurring microbes in algal microbiomes can be well explained by the latitudinal temperature gradient and associated break points in their beta diversity, with an average breakpoint at 14 °C ± 4.3, separating cold and warm upper oceans. As global warming impacts upper ocean temperatures, we project that break points of beta diversity move markedly pole-wards. Hence, abrupt regime shifts in algal microbiomes could be caused by anthropogenic climate change., Latitudinal ecosystem boundaries in the global upper ocean may be driven by many factors. Here the authors investigate pole-to-pole eukaryotic phytoplankton metatranscriptomes, gene co-expression networks, and beta diversity, finding that geographic patterns are best explained by temperature gradients.

The seaweed industry is growing worldwide to meet future resource needs in terms of food and fuel. In the meantime, the impact of expanding off-bottom seaweed cultivation on its environment is unclear. For example, it remains poorly understood how off-bottom seaweeds affect the local hydrodynamic environment, especially concerning turbulence that is more important for nutrient transport and availability than the mean flow velocity. Here, we carried out well-controlled flume experiments with mimic seaweed thalli, which are available, controllable, and stable, to investigate the impact of off-bottom seaweed canopies on whole-depth flow velocities in terms of both mean flow and turbulence velocity profiles. A careful comparison of behavior in the flow between natural and mimic seaweed thalli was made before these experiments. The results show that the floating seaweed thalli generate a surface boundary layer and have a profound impact on the velocity structure in the bottom boundary layer. More importantly, the generation, growth and dissipation of turbulence in the seaweed thalli area deeply affect the downstream distribution of near-bed turbulent strength and associated bed shear stress. Ignoring this turbulent variation would cause inaccurate predictions of morphological changes of the seabed. Our findings suggest that expanding the seaweed cultivation area may cause high risks of bed degradation and low diffusion in the downstream cultivation area. These findings provide novel insights into the environmental influence of off-bottom seaweed cultivation, with important implications for optimizing management strategies to promote seaweed productivity while minimizing seabed destabilization.

The microalgae Rhodomonas sp. is known as an excellent feed source for live feed organisms such as copepods. The main benefits of feeding Rhodomonas to live feed animals are attributed to the relative high polyunsaturated fatty acid (PUFA) level, the combination of containing both docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA), and the ratio between these fatty acids (FA). It has been shown that microalgae are able to accumulate valuable metabolites, such as lipids, under adverse conditions. The easiest and most inexpensive method to induce stress to microalgae is through nitrogen (N) starvation. In this study, the effect of N-starvation on biomass concentration, cell volume, and cellular composition, such as fatty acid concentration and composition, and phycoerythrin (PE) concentration of Rhodomonas sp. during a period of 8 days, was investigated. The research was divided into two stages. In the first (growth) stage, Rhodomonas sp. was cultivated in small 400 ml photobioreactors (Algaemist-S) under optimal conditions in turbidostat mode, which reached a biomass concentration of 1.5 gDW L–1 and dilution rate of 1.3 d–1. Samples were taken every 24 h for cell density and volume and productivity measurements in order to ensure a healthy and stable culture. In the next stage (N-starvation), the biomass was washed and transferred in a reactor filled with N-depleted medium. During N-starvation, samples were taken for biomass concentration, cell volume, PE and FA composition. The results of this study demonstrate that the lipid content increased significantly from 9% (t = 0 h) to 30% (t = 120 h) of the dry weight. After 120 h of N-starvation, the total FA content of Rhodomonas sp. remained stable for the remainder of the experiment (next 72 h). The highest increase of the FA concentration was represented by C16:0, C:18:1, C18:2, and C18:3, with highest concentrations after 120 h of starvation. The maximum EPA and DHA concentrations were observed after 48 h of starvation, while the maximum DHA to EPA ratio was detected at the end of the starvation.

Using either freshly pulped or preserved seaweed biomass for the extraction of protein can have a great effect on the amount of protein that can be extracted. In this study, the effect of four preservation techniques (frozen, freeze-dried, and air-dried at 40 and 70 °C) on the protein extractability, measured as Kjeldahl nitrogen, of four seaweed species, Chondrus crispus (Rhodophyceae), Ascophyllum nodosum, Saccharina latissima (both Phaeophyceae) and Ulva lactuca (Chlorophyceae), was tested and compared with extracting freshly pulped biomass. The effect of preservation is species dependent: in all four seaweed species, a different treatment resulted in the highest protein extractability. The pellet (i.e., the non-dissolved biomass after extraction) was also analyzed as in most cases the largest part of the initial protein ended up in the pellet and not in the supernatant. Of the four species tested, freeze-dried A. nodosum yielded the highest overall protein extractability of 59.6% with a significantly increased protein content compared with the sample before extraction. For C. crispus extracting biomass air-dried at 40 °C gave the best results with a protein extractability of 50.4%. Preservation had little effect on the protein extraction for S. latissima; only air-drying at 70 °C decreased the yield significantly. Over 70% of the initial protein ended up in the pellet for all U. lactuca extractions while increasing the protein content significantly. Extracting freshly pulped U. lactuca resulted in a 78% increase in protein content in the pellet while still containing 84.5% of the total initial total protein. These results show the importance of the right choice when selecting a preservation method and seaweed species for protein extraction. Besides the extracted protein fraction, the remaining pellet also has the potential as a source with an increased protein content.

Kelp life‐cycle transitions are complex and susceptible to various (a)biotic controls. Understanding the microscopic part of the kelp's lifecycle is of key importance, as gametophytes form a critical phase influencing, among others, the distributional limits of the species. Many environmental controls have been identified that affect kelp gametogenesis, whose interactive effects can be subtle and counterintuitive. Here we performed a fully factorial experiment on the (interactive) influences of light intensity, light quality, and the Initial Gametophyte Density (IGD) on Saccharina latissimi reproduction and vegetative growth of delayed gametophytes. A total of 144 cultures were followed over a period of 21 d. The IGD was a key determinant for reproductive success, with increased IGDs (≥0.04 mg DW · mL−1) practically halting reproduction. Interestingly, the effects of IGDs were not affected by nutrient availability, suggesting a resource‐independent effect of density on reproduction. The Photosynthetically Usable Radiation (PUR), overarching the quantitative contribution of both light intensity and light quality, correlated with both reproduction and vegetative growth. The PUR furthermore specifies that the contribution of light quality, as a lifecycle control, is a matter of absorbed photon flux instead of color signaling. We hypothesize that (i) the number of photons absorbed, independent of their specific wavelength, and (ii) IGD interactions, independent of nutrient availability, are major determinants of reproduction in S. latissimi gametophytes. These insights help understand kelp gametophyte development and dispersal under natural conditions, while also aiding the control of in vitro gametophyte cultures.

Through regular sampling surveys, the Flanders Marine Institute is generating long term data series for the Belgian coastal water and sand bank systems, a designated site in the Long Term Ecological Research (LTER) network. The data series is built on sampling activities initiated in 2002, but gradually upgraded and extended in the framework of the LifeWatch marine observatory and the Integrated Carbon Observation System (ICOS) participation. Nine nearshore stations are sampled monthly, with additional seasonal sampling of eight offshore stations. This paper presents the generated data series for nutrients, pigments, suspended matter and turbidity. The collection, methodology and processing of the 2002-2018 dataset is described, along with its data curation, integration and quality control. Yearly versions of the data are published online in a standardized format, accompanied with extensive metadata description and labelled with digital identifiers for traceability. Data is published under a CC-BY license, allowing use of the data under the condition of providing reference to the original source., SCOPUS: ar.j, info:eu-repo/semantics/published

Uptake rates of dissolved inorganic phosphorus and dissolved inorganic nitrogen under unsaturated and saturated conditions were studied in young sporophytes of the seaweeds Saccharina latissima and Laminaria digitata (Phaeophyceae) using a "pulse-and-chase" assay under fully controlled laboratory conditions. In a subsequent second "pulse-and-chase" assay, internal storage capacity (ISC) was calculated based on V

Climate change is affecting the hydrodynamics of the world’s oceans. How these changes will influence the productivity, distribution and abundance of phytoplankton communities is an urgent research question. Here we provide a unique high-resolution mesoscale description of the phytoplankton community composition in relation to vertical mixing conditions and other key physicochemical parameters along a meridional section of the Northeast Atlantic Ocean. Phytoplankton, assessed by a combination of flow cytometry and pigment fingerprinting (HPLC-CHEMTAX), and physicochemical data were collected from the top 250 m water column during the spring of 2011 and summer of 2009. Multivariate analysis identified water column stratification (based on 100 m depth-integrated Brunt-Vaisala frequency N2) as one of the key drivers for the distribution and separation of different phytoplankton taxa and size classes. Our results demonstrate that increased stratification (i) broadened the geographic range of Prochlorococcus as oligotrophic areas expanded northward, (ii) increased the contribution of picoeukaryotic phytoplankton to total autotrophic organic carbon (< 20 µm), and (iii) decreased the abundances of diatoms and cryptophytes. We discuss the implications of our findings for the classification of phytoplankton functional types in biogeochemical and ecological ocean models. As phytoplankton taxonomic composition and size affects productivity, biogeochemical cycling, ocean carbon storage and marine food web dynamics, the results provide essential information for models aimed at predicting future states of the ocean.

The oceans play a pivotal role in the global carbon cycle. It is not practical to measure the global daily production of organic carbon, the product of phytoplankton standing stock and its growth rate using discrete oceanographic methods. Instead, optical proxies from Earth-orbiting satellites must be used. To test the accuracy of optically derived proxies of phytoplankton physiology and growth rate, hyperspectral reflectance data from the wax and wane of a Phaeocystis bloom in laboratory mesocosms were compared with standard ex situ data. Chlorophyll biomass could be estimated accurately from reflectance using specific chlorophyll absorption algorithms. However, the conversion of chlorophyll (Chl) to carbon (C) was obscured by the non-linear increase in C : Chl under nutrient-limited growth. Although C : Chl was inversely correlated (r2 = 0.88) with the in situ fluorometric growth rate indicator Fv / Fm (Photosystem II quantum efficiency), none of them was linearly correlated to growth rate, constraining the accurate calculation of Phaeocystis growth or production rates. Unfortunately, the optical proxy &varphi;ph (quantum efficiency of fluorescence: the ratio of the number of fluoresced photons to the number of photons absorbed by the phytoplankton) did not show any correlation with Phaeocystis growth rate, and therefore it is concluded that &varphi;ph cannot be applied in the remotely sensed measurement of this species' carbon production rate.

Relationships between sea surface temperature (SST, > 10 m) and vertical density stratification, nutrient concentrations, and phytoplankton biomass, composition, and chlorophyll a (Chl a) specific absorption were assessed in spring and summer from latitudes 29 to 63° N in the northeast Atlantic Ocean. The goal of this study was to identify relationships between phytoplankton and abiotic factors in an existing SST and stratification gradient. Furthermore, a bio-optical model was used to estimate productivity for five phytoplankton groups. Nutrient concentration (integrated from 0 to 125 m) was inversely correlated with SST in spring and summer. SST was also inversely correlated with near-surface (0–50 m) Chl a and productivity for stratified stations. Near-surface Chl a and productivity showed exponential relationships with SST. Chl a specific absorption and excess light experiments indicated photoacclimation to lower irradiance in spring as compared to summer. In addition, Chl a specific absorption suggested that phytoplankton size decreased in summer. The contribution of cyanobacteria to water column productivity of stratified stations correlated positively with SST and inversely with nutrient concentration. This suggests that a rise in SST (over a 13–23 °C range) stimulates productivity by cyanobacteria at the expense of haptophytes, which showed an inverse relationship to SST. At higher latitudes, where rising SST may prolong the stratified season, haptophyte productivity may expand at the expense of diatom productivity. Depth-integrated Chl a (0–410 m) was greatest in the spring at higher latitudes, where stratification in the upper 200 m was weakest. This suggests that stronger stratification does not necessarily result in higher phytoplankton biomass standing stock in this region.

Economic and environmental developments in the last decades call for the displacement of fossil fuels to alternative energy sources. Biofuels are considered as a part of the solution for this challenge. Sustainable raw materials for the production of transportation biofuels such as biodiesel, biobutanol and bioethanol, can be obtained from algal biomass. In particular, marine macroalgal biomass is a promising feedstock for transportation biofuels because of (the) its fast growth and its potential cultivation on seawater, avoiding competition of resources with conventional agriculture of terrestrial plants used for food. In addition, dissolved inorganic nutrients like nitrogen, phosphorous and carbon are taken up by macroalgae, helping to alleviate eutrophication in seas and oceans. Using biological, chemical and engineering advances in the last decades, technologies to provide cost efficient cultivation, harvesting, extraction and processing of sustainable biofuels have to be elaborated. This paper provides a review of macroalgae based biorefineries with offshore cultivation and consequent biomass conversion into transportation liquid biofuels. We discuss the methods for offshore cultivation, harvesting, and conversion of macroalgae biomass into liquid transportation biofuels. Based on the current information and global experience, we present local perspectives specific for France, Germany, Norway, the Netherlands and Israel on the potential use of Exclusive Economic Zone for transportation biofuels production. Marketable suggestions for future research need to include all stakeholders of a given site for offshore biorefinery development.

The combined effects of different light and aqueous CO2 conditions were assessed for the Southern Ocean diatom Proboscia alata (Brightwell) Sundström in laboratory experiments. Selected culture conditions (light and CO2(aq) ) were representative for the natural ranges in the modern Southern Ocean. Light conditions were 40 (low) and 240 (high) μmol photons · m(-2) · s(-1) . The three CO2(aq) conditions ranged from 8 to 34 μmol · kg(-1) CO2(aq) (equivalent to a pCO2 from 137 to 598 μatm, respectively). Clear morphological changes were induced by these different CO2(aq) conditions. Cells in low [CO2(aq) ] formed spirals, while many cells in high [CO2(aq) ] disintegrated. Cell size and volume were significantly affected by the different CO2(aq) concentrations. Increasing CO2(aq) concentrations led to an increase in particulate organic carbon concentrations per cell in the high light cultures, with exactly the opposite happening in the low light cultures. However, other parameters measured were not influenced by the range of CO2(aq) treatments. This included growth rates, chlorophyll a concentration and photosynthetic yield (FV /FM ). Different light treatments had a large effect on nutrient uptake. High light conditions caused an increased nutrient uptake rate compared to cells grown in low light conditions. Light and CO2 conditions co-determined in various ways the response of P. alata to changing environmental conditions. Overall P. alata appeared to be well adapted to the natural variability in light availability and CO2(aq) concentration of the modern Southern Ocean. Nevertheless, our results showed that P. alata is susceptible to future changes in inorganic carbon concentrations in the Southern Ocean.

Phaeocystis globosa (Prymnesiophyceae) is an ecologically dominating phytoplankton species in many areas around the world. It plays an important role in both the global sulfur and carbon cycles, by the production of dimethylsulfide (DMS) and the drawdown of inorganic carbon. Phaeocystis globosa has a polymorphic life cycle and is considered to be a harmful algal bloom (HAB) forming species. All these aspects make this an interesting species to study the effects of increasing carbon dioxide (CO2) concentrations, due to anthropogenic carbon emissions. Here, the combined effects of three different dissolved carbon dioxide concentrations (CO2(aq)) (low: 4 μmol kg−1, intermediate: 6–10 μmol kg−1 and high CO2(aq): 21–24 μmol kg−1) and two different light intensities (low light, suboptimal: 80 μmol photons m−2 s−1 and high light, light saturated: 240 μmol photons m−2 s−1) are reported. The experiments demonstrated that the specific growth rate of P. globosa in the high light cultures decreased with increasing CO2(aq) from 1.4 to 1.1 d−1 in the low and high CO2 cultures, respectively. Concurrently, the photosynthetic efficiency (FV/FM) increased with increasing CO2(aq) from 0.56 to 0.66. The different light conditions affected photosynthetic efficiency and cellular chlorophyll a concentrations, both of which were lower in the high light cultures as compared to the low light cultures. These results suggest that in future inorganic carbon enriched oceans, P. globosa will become less competitive and feedback mechanisms to global change may decrease in strength.

The combined effects of different light and aqueous CO2 conditions were assessed for the Southern Ocean diatom Proboscia alata (Brightwell) Sundstrom in laboratory experiments. Selected culture conditions (light and CO2(aq)) were representative for the natural ranges in the modern Southern Ocean. Light conditions were 40 (low) and 240 (high) mu mol photons . m-2 . s-1. The three CO2(aq) conditions ranged from 8 to 34 mu mol . kg-1 CO2(aq) (equivalent to a pCO2 from 137 to 598 mu atm, respectively). Clear morphological changes were induced by these different CO2(aq) conditions. Cells in low [CO2(aq)] formed spirals, while many cells in high [CO2(aq)] disintegrated. Cell size and volume were significantly affected by the different CO2(aq) concentrations. Increasing CO2(aq) concentrations led to an increase in particulate organic carbon concentrations per cell in the high light cultures, with exactly the opposite happening in the low light cultures. However, other parameters measured were not influenced by the range of CO2(aq) treatments. This included growth rates, chlorophyll a concentration and photosynthetic yield (FV/FM). Different light treatments had a large effect on nutrient uptake. High light conditions caused an increased nutrient uptake rate compared to cells grown in low light conditions. Light and CO2 conditions co-determined in various ways the response of P. alata to changing environmental conditions. Overall P. alata appeared to be well adapted to the natural variability in light availability and CO2(aq) concentration of the modern Southern Ocean. Nevertheless, our results showed that P. alata is susceptible to future changes in inorganic carbon concentrations in the Southern Ocean.

The accuracy of remote sensing algorithms for phytoplankton biomass and physiology is difficult to test under natural conditions due to rapid changes in physical and biological forcings and the practical inability to manipulate nutrient conditions and phytoplankton composition in the sea. Therefore, an indoor mesocosm was designed to examine the optical properties of phytoplankton under controlled and manipulated conditions of irradiance, temperature, turbulence, and nutrient availability. Equipped with hyperspectral radiometers and bottom irradiance meters, it is shown that under semi-natural environmental conditions biogeochemically relevant species as Emiliania huxleyi and Phaeocystis globosa can be grown with good precision (+/- 20%) between duplicate mesocosms and between duplicate sensors (< 5% deviation). The accuracy of chlorophyll estimates by absorption, using an Integrating Cavity Absorption Meter, and fluorescence using water-leaving radiance was 74% to 80%, respectively, as it was negatively influenced by changes in phytoplankton physiology. Biomass detection was limitedto 1 to 2 mu g chlorophyll/L with an apparent linearity to 50 mu g chlorophyll/L. Estimates of the quantum efficiency of fluorescence (phi approximate to 0.01) were comparable to real-world estimates derived from satellite observations. It is concluded that the mesocosms adequately simulate natural conditions with sufficient accuracy and precision and that they offer an important tool in validating assumptions and hypotheses underlying remote sensing algorithms and models.

In the Eastern North Atlantic Ocean iron (Fe) speciation was investigated in three size fractions: the dissolvable from unfiltered samples, the dissolved fraction (o 0.2 mm) and the fraction smaller than 1000 kDa (o 1000 kDa). Fe concentrations were measured by flow injection analysis and the organic Fe complexation by voltammetry. In the research area the water column consisted of North Atlantic Central Water (NACW), below which Mediterranean Overflow Water (MOW) was found with the core between 800 and 1000 m depth. Below 2000 m depth the North Atlantic Deep Water (NADW) proper was recognised. Dissolved Fe and Fe in the o 1000 kDa fraction showed a nutrient like profile, depleted at the surface, increasing until 500–1000 m depth below which the concentration remained constant. Fe in unfiltered samples clearly showed the MOW with high concentrations (4 nM) compared to the overlying NACW and the underlying NADW, with 0.9 nM and 2 nM Fe, respectively. By using excess ligand (Excess L) concentrations as parameter we show a potential to bind Fe. The surface mixed layer had the highest excess ligand concentrations in all size fractions due to phytoplankton uptake and possible ligand production. The ratio of Excess L over Fe proved to be a complementary tool in revealing the relative saturation state of the ligands with Fe. In the whole water column, the organic ligands in the larger colloidal fraction (between 0.2 mm and 1000 kDa) were saturated with Fe, whereas those in the smallest fraction (o1000 kDa) were not saturated with Fe, confirming that this fraction was the most reactive one and regulates dissolution and colloid aggregation and scavenging processes. This regulation was remarkably stable with depth since the alpha factor (product of Excess L and K 0

Diatoms are the main primary producers in the Southern Ocean, governing the major nutrient cycles. Fragilariopsis kerguelensis (O’Meara) Hust. is the most abundant diatom species in the Southern Ocean and its paleo-oceanographic record is frequently used to reconstruct the past position and nutrient characteristics of the Antarctic polar front. Here we report on the responses of F. kerguelensis on prolonged exposure to a range of iron concentrations, allowing a characterization of morphological and nutrient-depletion changes in relation to iron status. Under iron limitation, F. kerguelensis grew slower, cells became smaller, chains became shorter, and the nutrient-depletion ratios changed. Prolonged exposure to iron limitation caused F. kerguelensis to decrease its surface area and volume 2-fold, and to increase its surface-to-volume ratio by 25%. With the decrease in growth rates, silicon (Si) and phosphorus (P) depletion per cell remained fairly constant, but when normalized per surface area (Si) or per cell volume (P), depletion increased. In contrast, nitrogen (N) depletion per cell decreased significantly together with the decrease in growth rates but was constant when normalized per cell volume. The different response in Si, P, and N depletion resulted in changes in the nutrient-depletion ratios, most notably in the Si:N ratio, which significantly increased, and in the N:P ratio, which significantly decreased with decreasing growth rates. It is concluded that under iron limitation, variation in cell size and/or nutrient depletion ultimately can cause changes in oceanic biogeochemical nutrient cycles. It enables the use of cell size of F. kerguelensis as a paleo-oceanographic proxy.

International audience; Vertical distributions of iron and iron binding ligands were determined in 2 size classes (dissolved < 0.2 μm, soluble < 200 kDa, e.g., ~ 0.03 μm) in the Southern Ocean. Colloidal iron and complexing capacity (> 200 kDa-< 0.2 μm) were inferred as the difference between the dissolved and soluble fractions. Dissolved iron and ligands exist primarily in the soluble size range in the surface waters, although iron-complexing colloids still represent a significant portion of the dissolved pool and this fraction increases markedly with depth. This work presents evidence for the colloidal nature of a significant portion (37-51% on average) of the 'dissolved' organic Fe pool in these oceanic waters. From the data it was not possible to discern whether iron colloids exist as discrete organic complexes and/or inorganic amorphous colloids. Iron-complexing colloids are the most saturated with iron at the thermodynamic equilibrium, whereas soluble organic ligands occur in larger excess compared to soluble iron. It suggests that the exchangeable fraction for iron uptake through dissociation of Fe complexes likely occurs in the soluble fraction, and that soluble ligands have the potential to buffer iron inputs to surface waters whereas iron colloids may aggregate and settle. Expectations based on Fe diffusion rates, distributions and the stability of the soluble iron complexes and iron colloids also suggest that the weaker soluble Fe complexes may be more bio-available, while the strongest colloids may be a major route for iron removal from oceanic waters. Investigations of the size classes of the dissolved organic iron thus can significantly increase our understanding of the oceanic iron cycle.

The major response to ocean iron fertilization is by large diatoms, which at Fe-replete ambient seawater show an optimum C:Fe elemental ratio of similar to 23 000 and a higher ratio of similar to 160 000 or more under Fe-limited conditions. The efficiency of CO2 drawdown during the several weeks of artificial fertilization experiments with concomitant observations is in the range of 100

Natural phytoplankton assemblages collected in surface waters above the Kerguelen Plateau or in the open-ocean and single-species cultures of Southern Ocean diatoms were used to address the existence and effects of natural iron fertilization near the Kerguelen Islands (Southern Ocean). The phytoplankton was transferred during so-called translocation experiments into water collected at the surface over the Plateau, open-ocean surface water or water collected close to the sediment of the Plateau. These watertypes differed in iron (iron-rich deep water and iron-poor surface water) and silicic acid concentration (silicic acid-rich Plateau deep and open-ocean surface water, silicic acid-poor Plateau surface water). As a general trend in the natural phytoplankton assemblages, cell numbers, chlorophyll autofluorescence, photosynthetic efficiency of photosystem II, chlorophyll a and phytoplankton carbon concentrations increased especially after translocation into Plateau deep water. This response was most pronounced in terms of increase in carbon assimilation in the larger-sized phytoplankton (>8 μm in cell diameter), mainly diatoms. Effects of translocation on bacteria and viruses followed those of the phytoplankton. Experiments with single-species cultures of large diatoms ( Fragilariopsis kerguelensis , Thalassiosira sp., Chaetoceros dichaeta ), which have high iron requirements, confirmed the observations made for the natural phytoplankton assemblages. Assuming a continuous flux of deep water to the surface over the Kerguelen Plateau, the translocation experiments provide evidence that this water contains the growth-stimulating factor, most likely iron, responsible for the formation of a phytoplankton bloom as is observed over the Kerguelen Plateau.

Short term (2 days) laboratory experiments were performed to study the change in irradiance induced production of Fe(II) in seawater in the presence of two open oceanic Southern Ocean diatom species, Thalassiosira sp. and Chaetoceros brevis. Three irradiance conditions were applied: 1) UVB+UVA+VIS, 2) UVA+ VIS, and 3) VIS, and Fe concentrations of 0 and 5 nM Fe were added to natural Southern Ocean seawater (containing 0.32 nM dissolved Fe and 1.69 equivalents of nM L(-1) Fe dissolved organic ligands, log K'=22.03). The photoproduced concentration of Fe(II) showed no relationship with the concentration of total dissolved Fe or the concentration of strongly chelated iron. During incubations with the diatoms an increase in the Fe(II) concentration during the second day suggested a modification of the Fe speciation. In the presence of Thalassiosira sp. photoreduction of Fe(III) was observed, whereas in the presence of C brevis irradiance independent Fe(III) reduction played an important role in the Fe(II) production. Furthermore, a decrease in the strongly chelated Fe concentration, in concert with a decrease in the conditional stability constant, suggested a modification of the strongly chelated Fe fraction in the experiments with C brevis. The chelated Fe fraction did not change in cultures with Thalassiosira sp. Overall, the presence of diatoms appeared to enhance the reactive Fe pool improving the biological availability of Fe. (c) 2007 Elsevier B.V. All rights reserved.

Shipboard and rernote-sensed Chlorophyll fluorescence were determined in the natural phytoplankton assemblage above the iron-enriched Kerguelen Plateau and the adjacent high-nutrient, low-Chlorophyll open Southern Ocean. The variance between fluorescence yield and photosynthetic efficiency was determined in combination with Chlorophyll a concentrations, irradiance and phytoplankton species distribution. A co-variance between the fluorescence measurements would allow the refinement of remote-sensing primary production algorithms. Distinct differences were found in photosynthetic efficiency and water-leaving fluorescence, with relatively high values for the Kerguelen Plateau and low values in the open ocean, reflecting the differences in Chlorophyll a concentrations. The co-variance of the fluorescence properties suggested that remote-sensed fluorescence measurements could be used to infer differences in the physiological state of the phytoplankton, hence primary production. Fluorescence yield, however, did not show the differences in the research area, most likely due to the low signal and the diurnal variation in water-leaving fluorescence.

Composition, physiology and growth response of 5 size classes of phytoplankton, rang- ing from 0.7 to 20 µm, was studied in an in situ iron enrichment experiment (EisenEx) and in bottle incubations in the Southern Ocean during austral spring 2000. In the field, iron enrichment resulted in only minor changes in numerical abundance, cell carbon content, photosynthetic efficiency (Fv:Fm) and the percentage of live cells of Synechococcus spp. and pico-eukaryotes (

The effects of light, silicate, or iron limitation on cell death rates and the accompanying effects on growth rates and photosynthetic efficiencies were investigated for 3 species of marine diatoms, Chaetoceros brevis, Chaetoceros calcitrans and Thalassiosira antarctica. Cell death rates were based on changes in time of the percentage dead cells using the SYTOX Green membrane per- meability assay. The death rates increased with decreasing irradiance levels and silicate concentra- tions for T. antarctica. In contrast, the cell death rates were hardly affected by low irradiances or sili- cate for C. brevis and C. calcitrans. Iron limitation experiments did not affect algal death rates significantly in the species tested. Growth rates under limiting conditions decreased from 0.6-0.4 to 0.2-0.1 d -1 , and only under low light conditions did growth halt completely. Photosynthetic efficien- cies did not always co-vary with cell death rates and were shown to be a sensitive indicator of light and iron limitation, but not of silicate limitation. This is the first detailed study providing data on diatom cell death under different growth-limiting conditions. The differential response in cell death rates indicates irradiance as a determining factor in diatom species succession and distribution.

International audience; Aerosol (soluble and total) iron and water-column dissolved (DFe, < 0.2 μm) and total dissolvable (TDFe, unfiltered) iron concentrations were determined in the Canary Basin and along a transect towards the Strait of Gibraltar, in order to sample across the Saharan dust plume. Cumulative dust deposition fluxes estimated from direct aerosol sampling during our one-month cruise are representative of the estimated deposition fluxes based on near surface water dissolved aluminium concentrations measured on board. Iron inventories in near surface waters combined with flux estimates confirmed the relatively short residence time of DFe in waters influenced by the Saharan dust plume (6–14 months). Enhanced near surface water concentrations of DFe (5.90–6.99 nM) were observed at the Strait of Gibraltar mainly due to inputs from metal-rich rivers. In the Canary Basin and the transect towards Gibraltar, DFe concentrations (0.07–0.76 nM) were typical of concentrations observed in the surface North Atlantic Waters, with the highest concentrations associated with higher atmospheric inputs in the Canary Basin. Depth profiles showed that DFe and TDFe were influenced by atmospheric inputs in this area with an accumulation of aeolian Fe in the surface waters. The sub-surface minimum of both DFe and TDFe suggests that a simple partitioning between dissolved and particulate Fe is not obvious there and that export may occur for both phases. At depths of around 1000–1300 m, both regeneration and Meddies may explain the observed maximum. Our data suggest that, in deep waters, higher particle concentrations likely due to dust storms may increase the scavenging flux and thus decrease DFe concentrations in deep waters.

Dissolved Fe and ligand concentrations and the Fe-binding strength of the organic ligands were measured in samples from the upper water column (150 m) of the oligotrophic waters of the Canary Basin (eastern North Atlantic Ocean). Concentrations of major nutrients, phytoplankton abundance and photosynthetic characteristics were also measured in the same samples.The concentrations of dissolved Fe and dissolved organic ligands were low with mean values of 0.31 +/- 0.18 nM Fe and 1.79 +/- 0.73 nEq of M Fe(n =47), respectively. The conditional binding constant varied between 10(19.8)-10(22.7) (n = 47). The largest variation with depth in the ligand concentrations (between 4.78 and 1.1 nEq of M Fe) was observed in the upper layer, above the Deep Chlorophyll Maximum (DCM located between 80 and 100 in), with high surface values in stations at 18 and 34.At the DCM where Fe was depleted, the ligand concentrations were still relatively high showing the same trend with depth as the amount of phytoplankton cells. Here 62% of the vertical variation in ligand concentrations can be explained by parameters describing phytoplankton cell abundance or biomass and orthosilicic acid concentration, which could reflect diatom growth. Ligand concentrations below the maximum of the DCM (n=4) showed good linear positive relationships with the total phytoplankton biomass as well as with 2 out of 4 distinguished groups of phytoplankton (Synechococcus and pico-eukaryote I).In the maximum of the DCM and below this maximum the phytoplankton origin of the dissolved organic ligands of Fe is very probable. Data suggest a release of ligands by cell lysis and not by an active production. However, the origin in the surface layer is more difficult to explain. Although the amount of phytoplankton cells in the surface layer is reduced, it is still similar to 25% of the cell concentration observed in the DCM. High concentrations of organic ligands could then be a remnant of past blooms or present production under nutrient depleted conditions. Input of Sahara dust can be another source of ligands. (c) 2006 Elsevier B.V. All rights reserved.

Diatoms in the Southern Ocean are limited by iron and light, and therefore produce little biomass. Sufficient biomass for analysis under these conditions requires large sample volumes, and diatom samples are therefore often pre-concentrated on a filter. A method for the digestion of diatom cells on polycarbonate filters, that is also suitable for trace metal analysis, is described here. Additional analysis by inductively coupled plasma-optical emission spectroscopy (ICP-OES) is used for the determination of biogenic silicon. Although several procedures were tested, the method of Hauptkorn et al., which uses tetramethylammonium hydroxide for the destruction of silicon is adapted here [Hauptkorn et al. (2001) Fres J Anal Chem 370:246–250]. Additional nitric acid is added to destroy the polycarbonate filters. The described method results in clear digests and a good correlation between cell numbers and silicon content. Using this procedure, the cellular silicon content for Chaetoceros brevis was determined as 86 ± 4 fmol cell−1. For Thalassiosira sp. a sensitivity effect was observed, and silicon content was determined as \( Si{\text{ = 4}}{\text{.2x10}}^{{{\text{ - 7}}}} {\left( {{\text{1 - exp}}^{{{\text{ - 2}}{\text{.2x10}}^{{{\text{ - 7}}}} {\text{cells}}}} } \right)} \). The obtained cellular silicon contents are in good agreement with values presented in the literature.

The determination of unidirectional fluxes of Fe in the seawater Fe-pools requires the use of Fe (radio)isotopes, which in turn necessitates off-line iron counting. To implement new off-line counting approaches, methods by King et al. (1991) [King DW, Lin J, Kester DR. Spectrophotometric determination of iron(II) in seawater at nanomolar concentrations. Anal Chim Acta 1991;247:125–32] were adapted for use with radio-iron for Fe(II) and total Fe analysis. The approach consists of passing samples through preloaded SepPak® C 18 cartridges with either ferrozine (FZ) for Fe(II) measurement or 1-nitroso-2-naphthol (1N2N) for total iron analysis. Cartridges are afterwards eluted with methanol and the eluate subsequently counted for radio-iron. A series of experiments was carried out to investigate the new method's applicability. This included loading and extraction behaviour, selectivity for the targeted iron species, iron species recovery, memory effects and the influence of iron concentration. The FZ-preloaded cartridge showed 85% Fe(II) recovery in seawater by direct determination. Recoveries of about 60% for Fe(II) in seawater were found when including 10–15 min waiting time and sample loops, which was probably caused by the instability of Fe(II) due to oxidation. Using 1N2N cartridges for total dissolved iron measurement, recovery was obtained as nearly 100%, but with high variability which was probably caused by the interaction time of Fe with the 1N2N-preloaded cartridges. The cartridges showed only small memory effects. Remaining radio-iron could only be removed from the cartridges with 0.5 M HCl, which destroyed the functionality of the cartridge. This result suggests that the Fe fraction, which could not be eluted by methanol, was irreversibly bound to the cartridge. Desferrioxamine B was applied as a model ligand to test for interference from naturally occurring organic ligands; the results indicate that interferences may occur and should be considered very carefully. The described approaches are tested for seawater iron concentrations in a 1.4–2.3 nM Fe range; further testing is necessary for other Fe concentration ranges.

Measurements of Fe(II) and H2O2 were carried out in the Atlantic sector of the Southern Ocean during EisenEx, an iron enrichment experiment. Iron was added on three separate occasions, approximately every 8 days, as a ferrous sulfate (FeSO4) solution. Vertical profiles of Fe(II) showed maxima consistent with the plume of the iron infusion. While H2O2 profiles revealed a corresponding minima showing the effect of oxidation of Fe(II) by H2O2, observations showed detectable Fe(II) concentrations existed for up to 8 days after an iron infusion. H2O2 concentrations increased at the depth of the chlorophyll maximum when iron concentrations returned to pre-infusion concentrations ( In this work, Fe(II) and dissolved iron were used as tracers themselves for subsequent iron infusions when no further SF6 was added. EisenEx was subject to periods of weak and strong mixing. Slow mixing after the second infusion allowed significant concentrations of Fe(II) and Fe to exist for several days. During this time, dissolved and total iron in the infusion plume behaved almost conservatively as it was trapped between a relict mixed layer and a new rain-induced mixed layer. Using dissolved iron, a value for the vertical diffusion coefficient Kz = 6.7±0.7 cm2 s−1 was obtained for this 2-day period. During a subsequent surface survey of the iron-enriched patch, elevated levels of Fe(II) were found in surface waters presumably from Fe(II) dissolved in the rainwater that was falling at this time. Model results suggest that the reaction between uncomplexed Fe(III) and O2− was a significant source of Fe(II) during EisenEx and helped to maintain high levels of Fe(II) in the water column. This phenomenon may occur in iron enrichment experiments when two conditions are met: (i) When Fe is added to a system already saturated with regard to organic complexation and (ii) when mixing processes are slow, thereby reducing the dispersion of iron into under-saturated waters.

An in situ iron enrichment experiment was carried out in the Southern Ocean Polar Frontal Zone and fertilized a patch of water within an eddy of the Antarctic Circumpolar Current (EisenEx, Nov. 2000). During the experiment, a physical speciation technique was used for iron analysis in order to understand the changes in iron distribution and size-fractionations, including soluble Fe ( 0.2 μm), throughout the development of the phytoplankton bloom. Prior to the first infusion of iron, dissolved (

International audience; Diatoms are a major component of phytoplankton community. They tend to dominate under natural high-nutrient concentrations, as well as during artificial Fe fertilisation experiments. They are main players in the biogeochemical cycle of carbon (C), as they can account for 40% of the total primary production in the Ocean and dominate export production, as well as in the biogeochemical cycles of the other macro-nutrients, nitrogen (N), phosphorus (P), and silicon (Si). Another important nutrient is Fe, which was shown to have a direct or indirect effect on nearly all the biogeochemical parameters of diatoms. In the present paper, an inventory is made of the growth, physiology and fate of many diatom species, including maximum growth rate, photosynthetic parameters (maximum specific rate of photosynthesis, photosynthetic efficiency and light adaptation parameter), nutrient limitation (half-saturation constant for growth/uptake), cellular elemental ratios, and loss terms (sinking rates, autolysis rates and grazing rates). This is a first step for improvement of the parameterisation of physiologically based phytoplankton growth and global 3D carbon models. This review is a synthesis of a large number of published laboratory experiments using monospecific cultures as well as field data. Our compilation confirms that size is an important factor explaining variations of biogeochemical parameters of diatoms (e.g. maximum growth rate, photosynthesis parameters, half-saturation constants, sinking rate, and grazing). Some variations of elemental ratios can be explained by adaptation of intracellular requirements or storage of Fe, and P, for instance. The important loss processes of diatoms pointed out by this synthesis are (i) sinking, as single cells as well as through aggregation which generally greatly increases sinking rate, (ii) cell autolysis, which can significantly reduce net growth rates, especially under nutrient limitation when gross growth rates are low, and (iii) grazing by both meso- and micro-zooplankton. This review also defines gaps concerning our knowledge on some important points. For example, we need to better know which iron species is available for phytoplankton, as well as the impact of Fe on the variation of the elemental ratios, especially in terms of assimilation and regeneration of C and N. A better quantification of prey selection by microzooplankton and mesozooplankton in natural environments is also needed, including preference for the various phytoplankton and zooplankton species as well as for aggregates and faecal pellets.

Picophytoplankton are a small or major component of the phytoplankton community and present in all oceanic systems, from pole to pole. They dominate in the low chlorophyll biomass areas, such as the (sub)tropical regions, but also contribute considerably (up to 20%) in the high chlorophyll biomass areas. The ecosystems of occurrence contrast significantly in physical and chemical settings. This includes a strongly mixed upper water column replete in nutrients as well as a strongly thermally stratified euphotic zone depleted in nutrients at the surface with a steep inverse light and nutrient gradient. These changes impose a strong impact on the composition of the picophytoplankton community but also on the biochemical and physiological properties of the species present. In particular, the pigmentation and cellular carbon, nitrogen and phosphorus quota and requirement will differ from a stratified compared to a well-mixed water column. As a result no characteristic values for the parameters required for this specific algal group in a global phytoplankton carbon model (the SWAMCO model,Lancelot et al. (2000), Deep-Sea Res. I, 47, 1621) can be given. In the present paper an inventory is made of the biochemical, physiological and photosynthetic parameters of two species of cyanobacteria (Prochlorococcus and Synechococcus) and the pico-size class fraction of the eukaryote phytoplankton component. Other groups of phytoplankton, such as diatoms, Trichodesmium, Phaeocystis and coccolithophorids, will be discussed in separate papers in this issue. This inventory is a mixture of laboratory experiments using well-defined algal populations as well as data derived from field surveys including a mixture of species. Where possible, the relevance of the parameters will be discussed in relation to the nature of the physico-chemical conditions of the area of occurrence.

The deposition of atmospheric dust into the ocean has varied considerably over geological time. Because some of the trace metals contained in dust are essential plant nutrients which can limit phytoplankton growth in parts of the ocean, it has been suggested that variations in dust supply to the surface ocean might influence primary production. Whereas the role of trace metal availability in photosynthetic carbon fixation has received considerable attention, its effect on biogenic calcification is virtually unknown. The production of both particulate organic carbon and calcium carbonate (CaCO3) drives the ocean's biological carbon pump. The ratio of particulate organic carbon to CaCO3 export, the so-called rain ratio, is one of the factors determining CO2 sequestration in the deep ocean. Here we investigate the influence of the essential trace metals iron and zinc on the prominent CaCO3-producing microalga Emiliania huxleyi. We show that whereas at low iron concentrations growth and calcification are equally reduced, low zinc concentrations result in a de-coupling of the two processes. Despite the reduced growth rate of zinc-limited cells, CaCO3 production rates per cell remain unaffected, thus leading to highly calcified cells. These results suggest that changes in dust deposition can affect biogenic calcification in oceanic regions characterized by trace metal limitation, with possible consequences for CO2 partitioning between the atmosphere and the ocean.

The atmosphere is widely recognized as a major source of Fe in the form of iron-containing dust. This study provides the first experiments in which the impact of dust on the growth rates of single species of Antarctic diatoms was assessed under laboratory conditions. The dust was among others characterized by x-ray powder diffraction analysis, analysis of total and amorphous Fe content, and dissolution rates of Fe in seawater. The amount of bioavailable Fe from the dust was determined, not via the complicated chemistry of Fe in seawater but by using diatoms as bioindicators for available Fe. Cultures of two large diatom species,Actinocyclussp. and Thalassiosira sp., were amended with potential dust aerosols from two dust-supplying regions, Namibia and Mauritania, and responses on growth rates were monitored. Apart from a difference in total Fe content, a difference in crystallinity existed in the Fe minerals of both dust types. The fraction of amorphous Fe was reflected in a higher reactivity/ dissolution of Fe in seawater. The increase in growth rate upon dust addition was positively related with the amount of amorphous Fe in the dust and with the dissolution rate of Fe in seawater. However, compared with equal FeCl3 concentrations, the dissolved Fe from the dust was not completely available for the diatoms. Interestingly, the diatoms used only a small part of the dissolved Fe, demonstrating the importance of algae as bioindicators.

Heme is the iron-containing prosthetic group of hemoproteins, and is thus required for photosynthesis, respiration and nitrate reduction in marine phytoplankton. Here we report concentrations of heme b in Southern Ocean phytoplankton and contrast our findings with those in coastal species. The concentration of particulate heme b (pmol l-1) observed at the end of the exponential growth phase was related to the concentration of dissolved iron in the culture media. Small Southern Ocean phytoplankton species (

The oceans play a pivotal role in the global carbon cycle. Unfortunately, the daily production of organic carbon, the product of phytoplankton standing stock and growth rate cannot be measured globally by discrete oceanographic methods. Instead, optical proxies from Earth-orbiting satellites must be used. To test the accuracy of optically-derived proxies of phytoplankton physiology and growth rate, standard ex situ data from the wax and wane of a Phaeocystis bloom in laboratory mesocosms were compared with hyperspectral reflectance data. Chlorophyll biomass could be estimated accurately from reflectance using specific chlorophyll absorption algorithms. However, the conversion of chlorophyll (Chl) to carbon (C) was obscured by the observed increase in C : Chl under nutrient-limited growth. C : Chl was inversely correlated (r2 = 0.88) with Photosystem II quantum efficiency (Fv/Fm), the in situ fluorometric oceanographic proxy for growth rate. In addition, the optical proxy for growth rate, the quantum efficiency of fluorescence &varphi; was linearly correlated to Fv/Fm (r2 = 0.84), but not – as by definition – by using total phytoplankton absorption, because during nutrient-limited growth the concentrations of non-fluorescent light-absorbing pigments increased. As a consequence, none of the three proxies (C : Chl, Fv/Fm, φ) was correlated to carbon or cellular phytoplankton growth rates. Therefore, it is concluded that although satellite derived estimates of chlorophyll biomass may be accurate, physiologically-induced non-linear shifts in growth rate proxies may obscure accurate phytoplankton growth rates and hence global carbon production estimates.

The interaction between iron and light limitation was investigated in 3 marine diatom species: 2 Antarctic diatom isolates, Chaetoceros brevis and C. dichaeta, and an isolate from temper- ate waters, C. calcitrans. In C. calcitrans and C. brevis, grown in the laboratory using trace metal buffered medium, both iron and light limitation affected growth rates, cellular chlorophyll a fluores- cence, spinal index (i.e., the number and size of spines) and cell size. Growth rates and cell size declined at lower iron concentrations and lower light intensities. Cellular chlorophyll a fluorescence increased with lower light, but decreased due to iron limitation. The spinal index, based on the ratio of side scatter to forward scatter, increased in iron- and light-limited cells. The large diatom C. dichaeta, grown in natural Southern Ocean water (without ethelenediaminetetraacetic acid (EDTA)) showed qualitatively similar responses to co-limitation by iron and light. C. dichaeta only grew under long-day light conditions. This response was further modified by the availability of iron. Addition of iron resulted in higher growth rates. In contrast, ambient iron concentrations did not limit the growth rate of the small Antarctic diatom C. brevis. However, iron limitation could be induced by addition of the natural iron binding ligand desferrioxamine B (DFOB). Addition of iron to Fe-depleted cultures of C. dichaeta and C. brevis reversed the effects of DFOB, as evidenced by rapid increases (within 24 h) in photochemical quantum efficiency (Fv/Fm) and decreases in effective absorption of the cross- section of photosystem II (σ PSII), the turnover time of the photosynthetic unit (τ) and the electron transfer rate (1/ τ), followed by an increase in growth rates after 48 h. The interactions between iron and light in Antarctic diatoms are sufficient to explain the observation that in the Southern Ocean some species (i.e., the small C. brevis) thrive under low iron and low light conditions, whereas other species (i.e., the large C. dichaeta) can bloom only under conditions of relatively high iron concentrations and favorable high light intensities or a long-day light period. These differences in physiological responses will have consequences for primary production, the carbon cycle and bio- geochemical cycles.

The remote waters of the Pacific region of the Southern Ocean are the furthest away from any upstream and upwind continental Fe sources. This prime area for expecting Fe limitation of the plankton ecosystem was studied (March-April 1995) along a north-south transect at similar to 89 degrees W. At the end of the austral summer the upper wind-mixed layers were in the order of similar to 100 m deep, thus mixing the algae down into the dimly lit part of the euphotic zone where photosynthesis is severely restricted. The dissolved Fe was found at low concentrations ranging from 0.05 nM near the surface to 0.5 nM in deeper waters. Along the transect (52 degrees S-69 degrees S), the dissolved iron was enhanced in the Polar Front, as well as near the Antarctic continental margin (0.6-1.0 nM). In between, the southern ACC branch was depleted with iron; here the concentrations in surface waters were quite uniform at about 0.21 nM. This is only somewhat lower than the 0.49 nM (October 1992) and 0.31 nM (November 1992) averages in early spring in the southern ACC part of Atlantic 6 degrees W sections [de Baar, H.J.W., de Jong, J.T.M., Bakker, D.C.E.. Loscher, B.M., Veth, C., Bathmann, U., Smetacek, V., 1995. Importance of iron for phytoplankton spring blooms and CO2 drawdown in the Southern Ocean. Nature 373, 412-415; Loscher, B.M., de Jong, J.T.M., de Baar, H.J.W., Veth, C., Dehairs, F., 1997. The distribution of iron in the Antarctic Circumpolar Current. Deep-Sea Research II 44, 143-188.]. First, the lower similar to 0.21 nM in March-April 1995 may partly be due to continuation of the seasonal trend where the phytoplankton growth, albeit modest, was removing Fe from the surface waters. Secondly, the 89 degrees W Pacific stations are further downstream continental or seafloor sources than the Atlantic 6 degrees W section. In the latter case, the ACC water had passed through the Drake Passage and also over the Sandwich Plateau. Indeed for Drake Passage, intermediate Fe concentrations have been reported by others. The generally somewhat lower surface water Fe at the ACC and PF at 89 degrees W is consistent with the distance from sources and the late summer. It also would explain the very low abundance of phytoplankton (Chl a) in the region and the conspicuous absence of plankton blooms. In the subAntarctic waters north of the Polar Front there are no diatoms, let alone diatom blooms, due to low availability of silicate. Thus, it appears the biological productivity is suppressed due to iron deficiency in combination with the severe seasonal effects of wind mixing on the light climate, as well as regional silicate limitation for diatoms. (C) 1999 Elsevier Science B.V. All rights reserved.

Concentrations of dissolved Ni, Cu, Zn, Cd and Pb were measured in water samples collected during a cruise with R.V Pelagia (29-6/14-7-1993) in the northern North Sea and N.E. Atlantic Ocean. At least six depths (0–90 m) were sampled with modified Go-Flo samplers from a rubber zodiac. In the study area, the first 25 m were well mixed and stratification occurred below this depth. The local bloom of Emiliania huxleyi hardly affected the trace metals concentration, except for some removal of Cd as seen from its correlation with nitrate. The mean dissolved concentrations were for Ni (3.66 nM), Cu (1.61 nM), Zn (4.5 nM), Cd (48 pM) and Pb (108 pM). These concentrations are among the lowest reported for the North Sea and are of similar magnitude to those found in the eastern North Atlantic at the same latitude. Zn was the only exception with values 10 times higher compared to those in the Atlantic Ocean, suggesting external inputs, mainly atmospheric and possibly from surrounding land masses. The observed ratio Zn:Cd in the North Sea and estuaries is in between the high ratio 600–900 for continental sources and the low ratio 5–10 for oceanic waters. Latter low ratio is consistent with the 21-fold stronger inorganic complexation of Cd in seawater which, in combination with the preferential biological uptake of Zn, may lead to the observed about hundredfold fractionation of Zn versus Cd in the marine system. Other processes may play a role but would need further investigation. The dissolved Pb values tend to be lower than found before in the North Sea, indicating decreasing inventories due to reduced anthropogenic emissions.