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As the annual expanse of Arctic summer ice‐cover steadily decreases, concomitant biogeochemical and ecological changes in this region are likely to occur. Because the Central Arctic Ocean is often nutrient and light limited, it is essential to understand how environmental changes will affect productivity, phytoplankton species composition, and ensuing changes in biogeochemistry in the region. During the transition from late summer to early autumn, water column sampling of various biogeochemical parameters was conducted along an ice‐floe drift station near the North Pole. Our results show that as the upper water column stratification weakened during the late summer–early autumn transition, nutrient concentrations, particulate dimethylsulfoniopropionate (DMSPp) levels, photosynthetic efficiency, and biological productivity, as estimated by ΔO2/Ar ratios, all decreased. Chemotaxonomic (CHEMTAX) analysis of phytoplankton pigments revealed a taxonomically diverse picoautotrophic community, with chlorophyll (Chl) c3‐containing flagellates and the prasinophyte, Pyramimonas spp., as the most abundant groups, comprising ~ 30% and 20% of the total Chl a (TChl a) biomass, respectively. In contrast to previous studies, the picoprasinophyte, Micromonas spp., represented only 5% to 10% of the TChl a biomass. Of the nine taxonomic groups identified, DMSPp was most closely associated with Pyramimonas spp., a Chl b‐containing species not usually considered a high DMSP producer. As the extent and duration of open, ice‐free waters in the Central Arctic Ocean progressively increases, we suggest that enhanced light transmission could potentially expand the ecological niche of Pyramimonas spp. in the region.

Marine phytoplankton play critical roles in the biogeochemistry of open and coastal oceans. However, the impact that individual species have on an ecosystem-wide scale can strongly depend on the production of cellular compounds, especially those that are climatically active such as dimethylsulfide (DMS). Herein, we use sorting flow cytometry to separate a distinct phytoplankton population from four samples taken along the Patagonian shelf near the Falkland Islands. Morphological, genetic, and biochemical analyses demonstrated that three of the sorted samples were dominated by a bloom of the dinoflagellate Prorocentrum minimum. Cellular quotas of the DMS-precursor dimethylsulfoniopropionate (DMSP) ranged from 1.23–4.11 pg cell−1 in the same population at different sampling stations. Causes of this variability may be due to different growth stages of the P. minimum bloom or changes in other environmental variables. Overall, in situ intracellular DMSP concentrations were lower than what would be expected based on previous, culture-based measurements. We demonstrate the difficulties inherent in sorting individual phytoplankton species from natural samples in order to determine in situ species-specific cellular quotas of important biogeochemical compounds.

Protein markers of cyanobacterial stress Nutrients including iron, nitrogen, and phosphorus limit primary productivity in the oceans. Determining how abundant cyanobacteria such as Prochlorococcus adapt to nutrient stress across marine settings requires accurate molecular assays. Saito et al. developed a proteomic and metaproteomic approach capable of targeting specific metabolic biomarkers from mixed communities in seawater (see the Perspective by Moore). Prochlorococcus proteins are indicative of a major limiting nutrient across a wide transect in the Pacific Ocean; however, they also show that the limitation of multiple nutrients at overlapping biomes is an additional source of stress. Science, this issue p. 1173 ; see also p. 1120

SummaryPhytoplankton blooms are ephemeral events of exceptionally high primary productivity that regulate the flux of carbon across marine food webs [1–3]. Quantification of bloom turnover [4] is limited by a fundamental difficulty to decouple between physical and biological processes as observed by ocean color satellite data. This limitation hinders the quantification of bloom demise and its regulation by biological processes [5, 6], which has important consequences on the efficiency of the biological pump of carbon to the deep ocean [7–9].Here, we address this challenge and quantify algal blooms’ turnover using a combination of satellite and in situ data, which allows identification of a relatively stable oceanic patch that is subject to little mixing with its surroundings. Using a newly developed multisatellite Lagrangian diagnostic, we decouple the contributions of physical and biological processes, allowing quantification of a complete life cycle of a mesoscale (∼10–100 km) bloom of coccolithophores in the North Atlantic, from exponential growth to its rapid demise. We estimate the amount of organic carbon produced during the bloom to be in the order of 24,000 tons, of which two-thirds were turned over within 1 week. Complimentary in situ measurements of the same patch area revealed high levels of specific viruses infecting coccolithophore cells, therefore pointing at the importance of viral infection as a possible mortality agent. Application of the newly developed satellite-based approaches opens the way for large-scale quantification of the impact of diverse environmental stresses on the fate of phytoplankton blooms and derived carbon in the ocean.

Over the last two decades, escalating rates of coastal development have altered macro- and micronutrient loading patterns to many southeastern U.S. estuaries. This study addresses a potential consequence of urbanization-associated coastal deforestation that may have important implications for estuarine ecosystem productivity: a reduction in bioavailable Fe. We compared the potential for Fe limitation at representative sites in two neighboring South Carolina salt marsh estuaries, one (“Parsonage Creek” site in Murrells Inlet estuary) impacted by urbanization-associated clear-cutting, and the other (“Oyster Landing” site in North Inlet estuary) undeveloped and surrounded by forests. The urbanized estuarine site was marked by lower dissolved Fe concentrations compared to the forested estuarine site. In bioassay experiments conducted from 1996 to 1999, the addition of chelated Fe to natural phytoplankton populations stimulated chlorophyll a concentrations in water from Parsonage Creek on 6 of 11 sampling dates. Chlorophyll responses in Oyster Landing water also were observed on two of those sampling times, dates that followed periods of relatively low rainfall. In one experiment, the addition of Fe+NO 3 − to Parsonage Creek water led to significantly higher chlorophyll concentrations than when NO 3 − was added alone, but Fe+NH 4 + addition did not result in higher chlorophyll than NH 4 + addition alone. It was hypothesized that the difference in NO 3 − and NH 4 + responses was related to a greater tendency towards Si depletion in Fe-stressed NO 3 − -grown samples, or alternatively, to the higher Fe requirement for NO 3 − assimilation relative to that needed for NH 4 + assimilation. In bioassay experiments using Fe-deplete semi-continuous cultures of Synechococcus WH8101, Fe addition did not affect the net growth rate of cultures transferred to water from either estuary, but increased the chlorophyll a content of cells transferred to Parsonage Creek water. Finally, photosynthesis vs. irradiance parameters ( α , P max ) obtained from Oyster Landing samples ( Synechococcus continuous cultures, while the same parameters derived from Parsonage Creek samples were much lower and comparable to those from Fe-deplete cultures. The results indicate that phytoplankton population growth from high salinity salt marsh estuaries can be Fe-limited. Also, the potential for Fe limitation was more prevalent in a urbanized deforested estuary than in an undeveloped forested estuary, consistent with the hypothesis that organically bound Fe from coastal forests plays an important role in supplying Fe for the growth of estuarine phytoplankton.

Phytoplankton pigments were measured using HPLC during non-ENSO conditions in mid-summer along a South Pacific transect from 67° S to the equator along 170° W. Highest concen- trations of chlorophyll a (chl a) occurred in the Polar and the Subtropical Fronts (PF and STF, respec- tively) with concentrations exceeding 500 ng l -1 . In the STF, there was a distinct subsurface chl a maximum (SCM) at 40 m, which gradually deepened northward to 120 m in the Subtropical Conver- gence Zone. Northwards, the SCM shoaled to about 30 m in the Equatorial Zone (EZ). Relatively high concentrations of fucoxanthin and 19'-butanoyloxyfucoxanthin occurred in the nutrient-rich waters south of the Subantarctic Front, and CHEMTAX analyses indicated that diatoms, chrysophytes, pelagophytes, and haptophytes dominated the phytoplankton assemblage. Northward of the PF to the STF, where silicate concentrations were

We report the results of a field incubation experiment demonstrating a substantial shift in the taxonomic composition of Equatorial Pacific phytoplankton assemblages exposed to CO2 levels of 150 and 750 ppm (dissolved CO2 ~3 to 25 µM). By the end of the experiment, the phytoplankton community in all samples was dominated by diatoms and Phaeocystis sp. However, the relative abun- dance of these phytoplankton taxa differed significantly between CO2 treatments. Taxonomic pig- ment analysis and direct microscopic examination of samples revealed that the abundance of diatoms decreased by ~50% at low CO2 relative to high CO2, while the abundance of Phaeocystis increased by ~60% at low CO2. This CO2-dependent shift was associated with a significant change in nutrient utilization, with higher ratios of nitrate:silicate (N:Si) and nitrate:phosphate (N:P) consumption by phytoplankton in the low CO2 treatment. Despite the significant changes in taxonomic composition and nutrient consumption ratios, total biomass and primary productivity did not differ significantly between the CO2 treatments. Our results suggest that CO2 concentrations could potentially influence competition among marine phytoplankton taxa and affect oceanic nutrient cycling.

Shipboard bottle incubation experiments were performed to investigate the effects of iron, light, and silicate on algal production of particulate dimethylsulfoniopropionate (DMSPp) in the Subantarctic Zone (SAZ) south of Tasmania during March 1998. Iron enrichment resulted in threefold to ninefold increases in DMSPp concentrations relative to control treatments, following 7 and 8-day incubation experiments. Additions of Fe and Si preferentially stimulated the growth of lightly-silicified pennate diatoms and siliceous haptophytes, respectively, to which we attribute the increased DMSPp production in the incubation bottles. Both of these algal groups were previously believed to be low DMSPp producers; however, our experimental data suggest that addition of iron and silicate to the low-silicate low-iron waters of the SAZ will result in increased production of DMSPp by lightly silicified diatoms and siliceous haptophytes, respectively. Increased irradiance enhanced DMSPp production in iron-amended treatments with both low (0.5 nM) and high (5 nM) concentrations of added iron. However, the role of light in stimulating DMSPp production was apparently of secondary importance compared to the effects of iron addition. The combination of high irradiance and high iron enrichment produced the highest DMSPp production in the experiments, suggesting that iron and light may have a synergistic effect in limiting algal DMSPp production in subantarctic waters.

The influence of irradiance and iron (Fe) supply on phytoplankton processes was investigated, north (47S, 142E) and south (54S, 142E) of the Subantarctic Front in austral autumn (March 1998). At both sites, resident cells exhibited nutrient stress (Fv/ Fm 0.3). Shipboard perturbation experiments examined two light (mean in situ and elevated) and two Fe (nominally 0.5 and 3 nM) treatments under silicic acid-replete conditions. Mean in situ light levels (derived from incident irradiances, mixed layer depths (MLDs), wind stress, and a published vertical mixing model) differed at the two sites, 25% of incident irradiance I 0 at 47S and 9% I 0 at 54S because of MLDs of 40 (47S) and 90 m (54S), when these stations were occupied. The greater MLD at 54S is reflected by tenfold higher cellular chlorophyll a levels in the resident phytoplankton. In the 47S experiment, chlorophyll a levels increased to 1 gL 1 only in the high-Fe treatments, regardless of irradiance levels, suggesting Fe limitation. This trend was also noted for cell abundances, silica production, and carbon fixation rates. In contrast, in the 54S experiment there were increases in chlorophyll a (to 2 gL 1 ), cell abundances, silica production, and carbon fixation only in the high-light treatments to which Fe had been added, suggesting that Fe and irradiance limit algal growth rates. Irradiance by altering algal Fe quotas is a key determinant of algal growth rate at 54S (when silicic acid levels are nonlimiting); however, because of the integral nature of Fe/light colimitation and the restricted nature of the current data set, it was not possible to ascertain the relative contributions of Fe and irradiance to the control of phytoplankton growth. On the basis of a climatology of summer mean MLD for subantarctic (SA) waters south of Australia the 47 and 54S sites appear to represent minimum and maximum MLDs, where Fe and Fe/ irradiance, respectively, may limit/colimit algal growth. The implications for changes in the factors limiting algal growth with season in SA waters are discussed.

Subantarctic Southern Ocean surface waters in the austral summer and autumn are characterized by high concentrations of nitrate and phosphate but low concentrations of dissolved iron (Fe, ∼0.05 nM) and silicic acid (Si

We investigated the hydrography, nutrients, and dissolved and particulate carbon pools in the western Pacific sector of the Antarctic Circumpolar Current (ACC) during austral summer 1996 to assess the region's role in the carbon cycle. Low fCO2 values along two transects indicated that much of the study area was a sink for atmospheric CO2. The fCO2 values were lowest near the Polar Front (PF) and the Subtropical Front (STF), concomitant with maxima of chlorophyll a and particulate and dissolved organic carbon. The largest biomass accumulations did not occur at fronts, which had high surface geostrophic velocities (20–51 cm s−1), but in relatively low velocity regions near fronts or in an eddy. Thus vertical motion and horizontal advection associated with fronts may have replenished nutrients in surface waters but also dispersed phytoplankton. Although surface waters north of the PF have been characterized as a “high nutrient-low chlorophyll” region, low silicic acid (Si) concentrations (2–4 μM) may limit production of large diatoms and therefore the potential carbon flux. Low concentrations (4–10 μM Si) at depths of winter mixing constrain the level of Si replenishment to surface waters. It has been suggested that an increase in aeolian iron north of the PF may increase primary productivity and carbon export. Our results, however, indicate that while diatom growth and carbon export may be enhanced, the extent ultimately would be limited by the vertical supply of Si. South of the PF, the primary mechanism by which carbon is exported to deep water appears to be through diatom flux. We suggest that north of the PF, particulate and dissolved carbon may be exported primarily to intermediate depths through subduction and diapycnal mixing associated with Subantarctic Mode Water and Antarctic Intermediate Water formation. These physical-biological interactions and Si dynamics should be included in future biogeochemical models to provide a more accurate prediction of carbon flux.

Dissolved iron and manganese and total dissolvable iron were measured in water column samples collected from the polynya region of the southern Ross Sea during cruises in November–December 1994 (spring 1994) and December 1995 to January 1996 (summer 1995). Iron and manganese addition bottle incubation experiments were also performed during these cruises in order to assess the nutritional sufficiency of ambient iron and manganese concentrations for growth of the phytoplankton community. Generally high dissolved iron concentrations (>0.5 nM) and relatively complex iron and manganese vertical profiles were obtained during the spring 1994 cruise, compared with the summer 1995 data. Dissolved iron concentrations in the upper water column averaged 1.0 nM during spring 1994 and 0.23 nM in summer 1995, excluding two stations where concentrations exceeding 1 nM are attributed to inputs from melting sea ice. The observed differences in the distribution of iron and manganese between spring 1994 and summer 1995 are attributed to seasonal decreases in the up welling of bottom waters and melting of sea ice, which supply these metals into the upper water column, combined with the cumulative removal of iron and manganese from the water column throughout the spring and summer, due to biological uptake, vertical export and scavenging by suspended and sinking particles. Results of the metal addition bottle incubation experiments indicate that ambient dissolved iron concentrations are adequate for phytoplankton growth requirements during the spring and early summer, when algal production is highest and Phaeocystis antarctica dominates the algal community, whereas low dissolved iron concentrations limit algal community growth later in the summer, except in the stratified, iron-enriched waters near melting sea ice, where diatoms are able to bloom. Our observations and the inferred seasonal distributions of P. antarctica and diatoms in these waters suggest that iron availability and vertical mixing (i.e., irradiance) exert the primary controls on phytoplankton growth and community structure in the southern Ross Sea during the spring and summer.

Patterns of nutrient utilization and primary productivity (PP) in late austral spring and early summer in the southwestern Ross Sea were characterized with respect to phytoplankton taxonomic composition, polynya dynamics, and upper ocean hydrography during the 1996–1997 oceanographic program Research on Ocean-Atmosphere Variability and Ecosystem Response in the Ross Sea. Phytoplankton biomass in the upper 150 m of the water column ranged from 40 to 540 mg chlorophyll a (Chl a) m−2, exceeding 200 mg Chl a m−2 everywhere except the extreme northern and eastern boundaries of the Ross Sea polynya. Diatom biomass was greatest in the shallow mixed layers of Terra Nova Bay, while the more deeply mixed waters of the Ross Sea polynya were dominated by Phaeocystis antarctica. Daily production computed from the disappearance of NO3 (1.14 g C m−2 d−1) and total dissolved inorganic carbon (TDIC, 1.29 g C m−2 d−1) is consistent with estimates made from an algorithm forced with satellite measurements of Chl a (1.25 g C m−2 d−1) and from measurements of 14C uptake (1.33 g C m−2 d−1). Phytoplankton PP in the Ross Sea averaged 100 g C m−2 yr−1 during 1996–1997. Despite the early formation of the Terra Nova Bay polynya the diatom bloom there did not reach its peak PP until middle to late January 1997 (most likely because of more intense wind mixing in November), ∼6 weeks after the P. antarctica bloom in the Ross Sea polynya had reached the same stage of development. From 70 to 100% of the C and N deficits in the upper 150 m could be accounted for by particulate organic matter, indicating that there had been little dissolved organic matter production or export of particulate material prior to our cruise. This suggests that early in the season, PP and zooplankton grazing are decoupled in the southwestern Ross Sea. The NO3∶PO4 disappearance ratio in waters dominated by P. antarctica (19.0±0.61) was significantly greater than in waters where diatoms were most common (9.52±0.33), and both were significantly different from the Redfield N∶P ratio of 16. Vertical profiles of TDIC suggest that P. antarctica took up 110% more CO2 per mole of PO4 removed than did diatoms, an important consideration for climate models that estimate C uptake from the removal of PO4.

The temporal and spatial patterns of phytoplankton biomass, productivity, and particulate matter composition in the Ross Sea were assessed during cruises in January 1990 and February 1992. Biomass and primary productivity in the southern Ross Sea were greatest during mid-January, with surface chlorophyll concentrations, particulate organic carbon levels, and integrated primary productivity averaging 4.9 μg L−1, 0.54 mg L−1 and 2.63 g C m−2 d−1, respectively. Comparable mean concentrations and rates for February were 1.1 μg L−1, 0.29 mg L−1, and 0.78 g C m−2 d−1 (decreases of 76, 46, and 70%, respectively), indicative of the scale of temporal changes. A distinct south-north transition also was observed both in productivity and phytoplankton biomass, with the lowest values occurring in the northern Ross Sea. East-west gradients in phytoplankton biomass and composition occurred within the southern Ross Sea. The areal productivity of the Ross Sea ranged from 0.15 to 2.85 g C m−2 d−1 and is among the highest found in the entire Antarctic. Carbon:chlorophyll ratios were uniformly high but were highest (150) in 1990 in the diatom-dominated western Ross Sea. Surface growth rates were modest, averaging less than 0.2 day−1 during both seasons. We hypothesize that the marked seasonality in the region provides an environment in which net growth rates, although slow, are maximized through low loss rates and which allows biomass to accumulate in the surface layer. Furthermore, the temporal variations are quantitatively similar to the observed spatial variations. Therefore the dominant determinant of phytoplankton biomass and productivity at any one point on the Ross Sea continental shelf is the stage of the seasonal growth cycle.

The distribution of phytoplankton biomass and pigments was determined in the Ross Sea during late austral summer 1992. Large gradients in biomass were noted both in the east-west and north-south direction, with maximum particulate matter concentrations occurring in the southwest portion of the Ross Sea. Two xanthophyll pigments dominated the profiles, fucoxanthin (indicative of diatoms) and 19′ hexanoyloxyfucoxanthin (hex) (indicative of the prymnesiophyte Phaeocystis antarctica). Fucoxanthin concentrations were greatest near the coast of Victoria Land and within the northern transect, whereas hex concentrations were highest offshore in the SE Ross Sea. Particulate carbon:chlorophyll a (POC:chl) ratios in diatom-dominated waters of the western Ross Sea were relatively high (210). The POC:chl ratio for Phaeocystis antarctica populations from the SE Ross Sea was substantially less (92) than the ratios observed in diatom-dominated waters. Pigment 14C labeling indicated that phytoplankton carbon in the diatom-dominated northern Ross Sea accounted for approximately 30% of the POC, with a phytoplankton C:chl ratio of 130. Short-term (24 hour) vertical fluxes of pigments ranged from 3 to 40 μg chl a m−2 d−1. Diatom-dominated regions had greater fluxes of phaeophorbides, suggesting that metazoans were the most important grazers at these locations. In contrast, the phaeophytin/total phaeopigment ratio was highest in waters dominated by Phaeocystis antarctica. The distribution of phytoplankton biomass and pigments revealed a spatially variable distribution of taxa, one which clearly has important consequences to food-web dynamics, biogeochemical cycles, and vertical flux patterns in the Ross Sea.

[1] The Ross Sea polynya is among the most productive regions in the Southern Ocean and may constitute a significant oceanic CO2 sink. Based on results from several field studies, this region has been considered seasonally iron limited, whereby a “winter reserve” of dissolved iron (dFe) is progressively depleted during the growing season to low concentrations (∼0.1 nM) that limit phytoplankton growth in the austral summer (December–February). Here we report new iron data for the Ross Sea polynya during austral summer 2005–2006 (27 December–22 January) and the following austral spring 2006 (16 November–3 December). The summer 2005–2006 data show generally low dFe concentrations in polynya surface waters (0.10 ± 0.05 nM in upper 40 m, n = 175), consistent with previous observations. Surprisingly, our spring 2006 data reveal similar low surface dFe concentrations in the polynya (0.06 ± 0.04 nM in upper 40 m, n = 69), in association with relatively high rates of primary production (∼170–260 mmol C m−2 d−1). These results indicate that the winter reserve dFe may be consumed relatively early in the growing season, such that polynya surface waters can become “iron limited” as early as November; i.e., the seasonal depletion of dFe is not necessarily gradual. Satellite observations reveal significant biomass accumulation in the polynya during summer 2006–2007, implying significant sources of “new” dFe to surface waters during this period. Possible sources of this new dFe include episodic vertical exchange, lateral advection, aerosol input, and reductive dissolution of particulate iron.

[1] We calculate net community production (NCP) during summer 2005–2006 and spring 2006 in the Ross Sea using multiple approaches to determine the magnitude and consistency of rates. Water column carbon and nutrient inventories and surface ocean O2/Ar data are compared to satellite-derived primary productivity (PP) estimates and 14C uptake experiments. In spring, NCP was related to stratification proximal to upper ocean fronts. In summer, the most intense C drawdown was in shallow mixed layers affected by ice melt; depth-integrated C drawdown, however, increased with mixing depth. ΔO2/Ar-based methods, relying on gas exchange reconstructions, underestimate NCP due to seasonal variations in surface ΔO2/Ar and NCP rates. Mixed layer ΔO2/Ar requires approximately 60 days to reach steady state, starting from early spring. Additionally, cold temperatures prolong the sensitivity of gas exchange reconstructions to past NCP variability. Complex vertical structure, in addition to the seasonal cycle, affects interpretations of surface-based observations, including those made from satellites. During both spring and summer, substantial fractions of NCP were below the mixed layer. Satellite-derived estimates tended to overestimate PP relative to 14C-based estimates, most severely in locations of stronger upper water column stratification. Biases notwithstanding, NCP-PP comparisons indicated that community respiration was of similar magnitude to NCP. We observed that a substantial portion of NCP remained as suspended particulate matter in the upper water column, demonstrating a lag between production and export. Resolving the dynamic physical processes that structure variance in NCP and its fate will enhance the understanding of the carbon cycling in highly productive Antarctic environments.

The Ross Sea is home to some of the largest phytoplankton blooms in the Southern Ocean. Primary production in this system has previously been shown to be iron limited in the summer and periodically iron and vitamin B(12) colimited. In this study, we examined trace metal limitation of biological activity in the Ross Sea in the austral spring and considered possible implications for vitamin B(12) nutrition. Bottle incubation experiments demonstrated that iron limited phytoplankton growth in the austral spring while B(12), cobalt, and zinc did not. This is the first demonstration of iron limitation in a Phaeocystis antarctica-dominated, early season Ross Sea phytoplankton community. The lack of B(12) limitation in this location is consistent with previous Ross Sea studies in the austral summer, wherein vitamin additions did not stimulate P. antarctica growth and B(12) was limiting only when bacterial abundance was low. Bottle incubation experiments and a bacterial regrowth experiment also revealed that iron addition directly enhanced bacterial growth. B(12) uptake measurements in natural water samples and in an iron fertilized bottle incubation demonstrated that bacteria serve not only as a source for vitamin B(12), but also as a significant sink, and that iron additions enhanced B(12) uptake rates in phytoplankton but not bacteria. Additionally, vitamin uptake rates did not become saturated upon the addition of up to 95 pM B(12). A rapid B(12) uptake rate was observed after 13 min, which then decreased to a slower constant uptake rate over the next 52 h. Results from this study highlight the importance of iron availability in limiting early season Ross Sea phytoplankton growth and suggest that rates of vitamin B(12) production and consumption may be impacted by iron availability.

We investigated biological Fe recycling in plankton communities from two contrasting marine ecosystems, the equatorial Pacific and Monterey Bay, California. Our experiments involved addition of cultured cyanobacteria and diatoms labeled with Fe radiotracer to natural plankton communities. Transfer of label into the natural community was followed by size fractionation and microautoradiography, and intracellular Fe was determined with titanium wash techniques. We documented the transfer of labeled Fe from added cyanobacteria to indigenous large cells in both environments and from large diatoms to indigenous small cells in the equatorial Pacific. Microautoradiography and Ti washing verified that transfer occurred from and into intracellular fractions in Monterey Bay experiments, although substantial amounts of transferred Fe were also found to be surface-adsorbed or associated with detritus. We argue that, as is the case for N, recycling of cellular Fe to support regenerated production could be an important component of marine ecosystem dynamics, especially in areas low in Fe.

International audience; The CLAW hypothesis argues that a negative feedback mechanism involving phytoplankton-derived dimethylsulfoniopropionate (DMSP) could mitigate increasing sea surface temperatures that result from global warming. DMSP is converted to the climatically active dimethylsulfide (DMS), which is transferred to the atmosphere and photochemically oxidized to sulfate aerosols, leading to increases in planetary albedo and cooling of the Earth's atmosphere. A shipboard incubation experiment was conducted to investigate the effects of increased temperature and pCO2 on the algal community structure of the North Atlantic spring bloom and their subsequent impact on particulate and dissolved DMSP concentrations (DMSPp and DMSPd). Under 'greenhouse' conditions (elevated pCO2; 690 ppm) and elevated temperature (ambient + 4°C), coccolithophorid and pelagophyte abundances were significantly higher than under control conditions (390 ppm CO2 and ambient temperature). This shift in phytoplankton community structure also resulted in an increase in DMSPp concentrations and DMSPp:chl a ratios. There were also increases in DMSP-lyase activity and biomass-normalized DMSP-lyase activity under 'greenhouse' conditions. Concentrations of DMSPd decreased in the 'greenhouse' treatment relative to the control. This decline is thought to be partly due to changes in the microzooplankton community structure and decreased grazing pressure under 'greenhouse' conditions. The increases in DMSPp in the high temperature and greenhouse treatments support the CLAW hypothesis; the declines in DMSPd do not.

Environmental context Dimethylsulfoniopropionate (DMSP), a small sulfur compound biosynthesised by algae, plays an important role in global climate, particularly in polar regions. We investigated salinity effects on DMSP levels, and provide the first experimental measurements of DMSP and associated physiological changes in a polar diatom across to a range of gradual salinity shifts representative of sea-ice conditions. Quantitative estimates of DMSP in polar diatoms following salinity changes will facilitate new mathematical models to predict seasonal responses and reactions to climate change. Abstract Although extreme environmental gradients within sea-ice have been proposed to stimulate dimethylsulfoniopropionate (DMSP) accumulation in diatoms, a taxa whose temperate counterparts show relatively low concentrations, this has yet to be experimentally validated across a range of salinities representative of sea-ice conditions. The present study examined changes in DMSP concentrations in the widespread polar diatom Fragilariopsis cylindrus in response to gradual salinity shifts representative of those encountered during sea-ice formation and melt. DMSP concentrations were elevated up to 127% in 70-salinity cultures. Low-salinity shifts decreased intracellular DMSP concentrations in a gradient-dependent manner that suggests DMSP recycling rather than release under milder hyposalinity shifts. Permeable membranes were detected in ~45% of 10-salinity cells; therefore, loss of membrane integrity may only partially explain DMSP release in the lowest-salinity group. Growth rates, photosynthetic efficiency of photosystem II and reactive oxygen species detection indicated only partial impairment by salinity stress in this organism. Thus, experimental evidence supports the role of DMSP as a compatible solute in the acclimation of a sea-ice diatom across large salinity gradients and measurements of associated physiological changes will improve interpretation of environmental measurements.

International audience; Global climate change is predicted to have large effects on the ocean that could cause shifts in current algal community structure, major nutrient cycles, and carbon export. The Bering Sea is already experiencing changes in sea surface temperature (SST), unprecedented algal blooms, and alterations to trophic level dynamics. We incubated phytoplankton communities from 2 Bering Sea regimes under conditions of elevated SST and/or partial pressure of carbon dioxide (pCO2) similar to predicted values for 2100. In our 'greenhouse ocean' simulations, maximum biomass-normalized photosynthetic rates increased 2.6 to 3.5 times and community composition shifted away from diatoms and towards nanophytoplankton. These changes were driven largely by elevated temperature, with secondary effects from increased pCO2. If these results are indicative of future climate responses, community shifts towards nanophytoplankton dominance could reduce the ability of the Bering Sea to maintain the productive diatom-based food webs that currently support one of the world's most productive fisheries.

Annual Emiliania huxleyi blooms (along with other coccolithophorid species) play important roles in the global carbon and sulfur cycles. E. huxleyi blooms are routinely terminated by large, host-specific dsDNA viruses, (Emiliania huxleyi Viruses; EhVs), making these host-virus interactions a driving force behind their potential impact on global biogeochemical cycles. Given projected increases in sea surface temperature due to climate change, it is imperative to understand the effects of temperature on E. huxleyi's susceptibility to viral infection and its production of climatically active dimethylated sulfur species (DSS). Here we demonstrate that a 3°C increase in temperature induces EhV-resistant phenotypes in three E. huxleyi strains and that successful virus infection impacts DSS pool sizes. We also examined cellular polar lipids, given their documented roles in regulating host-virus interactions in this system, and propose that alterations to membrane-bound surface receptors are responsible for the observed temperature-induced resistance. Our findings have potential implications for global biogeochemical cycles in a warming climate and for deciphering the particular mechanism(s) by which some E. huxleyi strains exhibit viral resistance.

Iron availability and temperature are important limiting factors for the biota in many areas of the world ocean, and both have been predicted to change in future climate scenarios. However, the impacts of combined changes in these two key factors on microbial trophic dynamics and nutrient cycling are unknown. We examined the relative effects of iron addition (+1 nM) and increased temperature (+4°C) on plankton assemblages of the Ross Sea, Antarctica, a region characterized by annual algal blooms and an active microbial community. Increased iron and temperature individually had consistently significant but relatively minor positive effects on total phytoplankton abundance, phytoplankton and microzooplankton community composition, as well as photosynthetic parameters and nutrient drawdown. Unexpectedly, increased iron had a consistently negative impact on microzooplankton abundance, most likely a secondary response to changes in phytoplankton community composition. When iron and temperature were increased in concert, the resulting interactive effects were greatly magnified. This synergy between iron and temperature increases would not have been predictable by examining the effects of each variable individually. Our results suggest the possibility that if iron availability increases under future climate regimes, the impacts of predicted temperature increases on plankton assemblages in polar regions could be significantly enhanced. Such synergistic and antagonistic interactions between individual climate change variables highlight the importance of multivariate studies for marine global change experiments.

A new alga-bacterium model system sheds light on mortality factors affecting phytoplankton bloom demise and sulfur cycling., Emiliania huxleyi is a bloom-forming microalga that affects the global sulfur cycle by producing large amounts of dimethylsulfoniopropionate (DMSP) and its volatile metabolic product dimethyl sulfide. Top-down regulation of E. huxleyi blooms has been attributed to viruses and grazers; however, the possible involvement of algicidal bacteria in bloom demise has remained elusive. We demonstrate that a Roseobacter strain, Sulfitobacter D7, that we isolated from a North Atlantic E. huxleyi bloom, exhibited algicidal effects against E. huxleyi upon coculturing. Both the alga and the bacterium were found to co-occur during a natural E. huxleyi bloom, therefore establishing this host-pathogen system as an attractive, ecologically relevant model for studying algal-bacterial interactions in the oceans. During interaction, Sulfitobacter D7 consumed and metabolized algal DMSP to produce high amounts of methanethiol, an alternative product of DMSP catabolism. We revealed a unique strain-specific response, in which E. huxleyi strains that exuded higher amounts of DMSP were more susceptible to Sulfitobacter D7 infection. Intriguingly, exogenous application of DMSP enhanced bacterial virulence and induced susceptibility in an algal strain typically resistant to the bacterial pathogen. This enhanced virulence was highly specific to DMSP compared to addition of propionate and glycerol which had no effect on bacterial virulence. We propose a novel function for DMSP, in addition to its central role in mutualistic interactions among marine organisms, as a mediator of bacterial virulence that may regulate E. huxleyi blooms.