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Genomes reveal nutrient stress patterns Within the surface ocean, nitrogen, iron, and phosphorous can all be limiting nutrients for phytoplankton depending on location. Ustick et al. used the prevalence of Prochlorococcus genes involved in nutrient acquisition to develop maps of inferred nutrient stress across the global ocean (see the Perspective by Coleman). They found broad patterns of limitation consistent with an Earth system model and nutrient addition experiments. Leveraging metagenomic data in this manner is an appealing approach that will help to expand our understanding of the biogeochemistry in the vast open ocean. Science , this issue p. 287 ; see also p. 239

There is limited biogeochemical rate data from the oligotrophic central Indian Ocean, but it is known that there are geographical gradients in the physical and chemical conditions that may lead to unique biogeochemical regimes. As participants on a GO-SHIP repeat hydrography cruise, a transect was completed in spring of 2016 from 28 °S to 18 °N in the Indian Ocean, roughly along the 95 °E meridion. Cell count samples (phytoplankton and heterotrophic bacteria) analyzed by flow cytometry, and samples for carbon and nitrogen productivity incubations, assessed by stable isotopic tracers, were obtained from 20 m at approximately every other degree of latitude. Microbial cell counts by flow cytometry indicate that Prochlorococcus was the principal autotroph, but with increasing contributions of Synechococcus around the equator. Large eukaryotes (> 20 μm), imaged using FlowCAM, were generally absent or in very low abundance (

Variation in ocean C:N:P of particulate organic matter (POM) has led to competing hypotheses for the underlying drivers. Each hypothesis predicts C:N:P equally well due to regional co-variance in environmental conditions and biodiversity. The Indian Ocean offers a unique positive temperature and nutrient supply relationship to test these hypotheses. Here we show how elemental concentrations and ratios vary over daily and regional scales. POM concentrations were lowest in the southern gyre, elevated across the equator, and peaked in the Bay of Bengal. Elemental ratios were highest in the gyre, but approached Redfield proportions northwards. As Prochlorococcus dominated the phytoplankton community, biodiversity changes could not explain the elemental variation. Instead, our data supports the nutrient supply hypothesis. Finally, gyre dissolved iron concentrations suggest extensive iron stress, leading to depressed ratios compared to other gyres. We propose a model whereby differences in iron supply and N2-fixation influence C:N:P levels across ocean gyres., The Indian Ocean provides a unique environmental gradient to test underlying drivers of the elemental composition of particulate organic matter. Here the authors show that nutrient supply, over temperature and biodiversity changes, controls regional variation of elemental ratios in the tropical Indian Ocean.

The intertidal mussel Mytilus californianus is a critical foundation species that is exposed to fluctuations in the environment along tidal- and wave-exposure gradients. We investigated feeding and digestion in mussels under laboratory conditions and across environmental gradients in the field. We assessed whether mussels adopt a rate-maximization (higher ingestion and lower assimilation) or a yield-maximization acquisition (lower ingestion and higher assimilation) strategy under laboratory conditions by measuring feeding physiology and digestive enzyme activities. We used digestive enzyme activity to define resource acquisition strategies in laboratory studies, then measured digestive enzyme activities in three microhabitats at the extreme ends of the tidal- and wave-exposure gradients within a stretch of shore (, Summary: We show for the first time that the mussel Mytilus californianus has a flexible digestive strategy to deal with varying environmental conditions that are based upon their position on the shore.

The factors that control elemental ratios within phytoplankton, like carbon:nitrogen:phosphorus (C:N:P), are key to biogeochemical cycles. Previous studies have identified relationships between nutrient-limited growth and elemental ratios in large eukaryotes, but little is known about these interactions in small marine phytoplankton like the globally important Cyanobacteria. To improve our understanding of these interactions in picophytoplankton, we asked how cellular elemental stoichiometry varies as a function of steady-state, N- and P-limited growth in laboratory chemostat cultures of Synechococcus WH8102. By combining empirical data and theoretical modeling, we identified a previously unrecognized factor (growth-dependent variability in cell size) that controls the relationship between nutrient-limited growth and cellular elemental stoichiometry. To predict the cellular elemental stoichiometry of phytoplankton, previous theoretical models rely on the traditional Droop model, which purports that the acquisition of a single limiting nutrient suffices to explain the relationship between a cellular nutrient quota and growth rate. Our study, however, indicates that growth-dependent changes in cell size have an important role in regulating cell nutrient quotas. This key ingredient, along with nutrient-uptake protein regulation, enables our model to predict the cellular elemental stoichiometry of Synechococcus across a range of nutrient-limited conditions. Our analysis also adds to the growth rate hypothesis, suggesting that P-rich biomolecules other than nucleic acids are important drivers of stoichiometric variability in Synechococcus. Lastly, by comparing our data with field observations, our study has important ecological relevance as it provides a framework for understanding and predicting elemental ratios in ocean regions where small phytoplankton like Synechococcus dominates.

Current hypotheses suggest that cellular elemental stoichiometry of marine eukaryotic phytoplankton such as the ratios of cellular carbon:nitrogen:phosphorus (C:N:P) vary between phylogenetic groups. To investigate how phylogenetic structure, cell volume, growth rate, and temperature interact to affect the cellular elemental stoichiometry of marine eukaryotic phytoplankton, we examined the C:N:P composition in 30 isolates across 7 classes of marine phytoplankton that were grown with a sufficient supply of nutrients and nitrate as the nitrogen source. The isolates covered a wide range in cell volume (5 orders of magnitude), growth rate (

Journal of Plankton Research plankt.oxfordjournals.org J. Plankton Res. (2016) 38(4): 1052–1061. First published online January 18, 2016 doi:10.1093/plankt/fbv120 Contribution to the Themed Section: Scaling from individual p lankton to marine ecosystems Diel variability in the elemental composition of the marine cyanobacterium Synechococcus JOHANN S. LOPEZ 1 , NATHAN S. GARCIA 1 *, DAVID TALMY 2 AND ADAM C. MARTINY 1,3 DEPARTMENT OF EARTH SYSTEM SCIENCE , UNIVERSITY OF CALIFORNIA , IRVINE , CA , USA , DEPARTMENT OF EARTH , ATMOSPHERE AND PLANETARY SCIENCES , MASSACHUSETTS INSTITUTE OF TECHNOLOGY , CAMBRIDGE , MA , USA AND DEPARTMENT OF ECOLOGY AND EVOLUTIONARY BIOLOGY , UNIVERSITY OF CALIFORNIA , IRVINE , CA , USA * CORRESPONDING AUTHOR : n8garcia@gmail.com Received August 26, 2015; accepted December 10, 2015 Corresponding editor: Pia Moisander The ratio of elements such as carbon:nitrogen:phosphorus (C:N:P) in phytoplankton is known to vary substantially within single isolates and across environmental gradients. In addition, C:N:P is known to vary throughout the day due to diel patterns in nutrient acquisition and storage. It has been hypothesized that small phytoplankton such as marine cyanobacteria have relatively invariable elemental ratios during a 24 h period, whereas larger phytoplankton have a greater capacity to store elements and thus a wider diel range of C:N:P. To test this hypothesis, we examined diel vari- ability in cellular C:N:P, using a chemostat culturing system, for one of the most abundant marine cyanobacteria, Synechococcus (WH8102) during two 24 h periods. The cellular C quota nearly doubled during the 14 h light period and was subsequently reduced during the dark period. The cellular N quota also varied considerably, whereas the P quota remained relatively stable. These daily changes in elemental quotas led to highly variable C:N cell and C:P cell . Furthermore, the magnitude of variability in cellular elemental stoichiometry of Synechococcus was positively related to the growth rate. We constructed a model to test the extent to which variation in C:N cell and C:P cell is related to reserve carbon accumulation and depletion over each light –dark cycle. Results imply that, in addition to growth-related respiratory losses, Synechococcus also purges excess C during the dark period in order to maintain a nutritive balance within cells. Our data suggest that diel variation in C:N cell and C:P cell of Synechococcus is of the same order of magnitude as stoichiometric variation within plankton communities between major ocean environments. KEYWORDS: cyanobacteria; Synechococcus; stoichiometry; diel; Redfield; phytoplankton available online at www.plankt.oxfordjournals.org # The Author 2016. Published by Oxford University Press. All rights reserved. For permissions, please email: journals.permissions@oup.com

Marine dinitrogen (N2)-fixing cyanobacteria have large impacts on global biogeochemistry as they fix carbon dioxide (CO2) and fertilize oligotrophic ocean waters with new nitrogen. Iron (Fe) and phosphorus (P) are the two most important limiting nutrients for marine biological N2 fixation, and their availabilities vary between major ocean basins and regions. A long-standing question concerns the ability of two globally dominant N2-fixing cyanobacteria, unicellular Crocosphaera and filamentous Trichodesmium, to maintain relatively high N2-fixation rates in these regimes where both Fe and P are typically scarce. We show that under P-deficient conditions, cultures of these two cyanobacteria are able to grow and fix N2 faster when Fe deficient than when Fe replete. In addition, growth affinities relative to P increase while minimum concentrations of P that support growth decrease at low Fe concentrations. In Crocosphaera, this effect is accompanied by a reduction in cell sizes and elemental quotas. Relatively high growth rates of these two biogeochemically critical cyanobacteria in low-P, low-Fe environments such as those that characterize much of the oligotrophic ocean challenge the common assumption that low Fe levels can have only negative effects on marine primary producers. The closely interdependent influence of Fe and P on N2-fixing cyanobacteria suggests that even subtle shifts in their supply ratio in the past, present and future oceans could have large consequences for global carbon and nitrogen cycles.

We examined the combined effects of light and pCO2 on growth, CO2-fixation and N2-fixation rates by strains of the unicellular marine N2-fixing cyanobacterium Crocosphaera watsonii with small (WH0401) and large (WH0402) cells that were isolated from the western tropical Atlantic Ocean. In low-pCO2-acclimated cultures (190 ppm) of WH0401, growth, CO2-fixation and N2-fixation rates were significantly lower than those in cultures acclimated to higher (present-day ∼385 ppm, or future ∼750 ppm) pCO2 treatments. Growth rates were not significantly different, however, in low-pCO2-acclimated cultures of WH0402 in comparison with higher pCO2 treatments. Unlike previous reports for C. watsonii (strain WH8501), N2-fixation rates did not increase further in cultures of WH0401 or WH0402 when acclimated to 750 ppm relative to those maintained at present-day pCO2. Both light and pCO2 had a significant negative effect on gross : net N2-fixation rates in WH0402 and trends were similar in WH0401, implying that retention of...

The diazotrophic cyanobacteria Trichodesmium spp. contribute approximately half of the known marine dinitrogen (N2 ) fixation. Rapidly changing environmental factors such as the rising atmospheric partial pressure of carbon dioxide (pCO2 ) and shallower mixed layers (higher light intensities) are likely to affect N2 -fixation rates in the future ocean. Several studies have documented that N2 fixation in laboratory cultures of T. erythraeum increased when pCO2 was doubled from present-day atmospheric concentrations (∼380 ppm) to projected future levels (∼750 ppm). We examined the interactive effects of light and pCO2 on two strains of T. erythraeum Ehrenb. (GBRTRLI101 and IMS101) in laboratory semicontinuous cultures. Elevated pCO2 stimulated gross N2 -fixation rates in cultures growing at 38 μmol quanta · m(-2 ) · s(-1) (GBRTRLI101 and IMS101) and 100 μmol quanta · m(-2 ) · s(-1) (IMS101), but this effect was reduced in both strains growing at 220 μmol quanta · m(-2 ) · s(-1) . Conversely, CO2 -fixation rates increased significantly (P

We demonstrated that the toxicity of the harmful bloom dinoflagellate Karlodinium veneficum is regulated by both CO 2 concentrations and phosphate availability. Semi-continuous cultures were grown in a factorial experiment under all combinations of 3 CO 2 levels (230, 430 and 745 ppm) and 2 phosphate conditions (0.5 and 20 uM). After steady-state acclimation was achieved, karlotoxin cellular quotas and growth rates were determined in all 6 treatments. This strain produced both types of karlotoxin, KmTx-1 and KmTx-2. Chlorophyll a-normalized production of the 2 types of karlotoxins was much higher in P-limited cultures compared with P-replete ones under the same CO 2 conditions. Increasing CO 2 strongly stimulated production of KmTx-1 and decreased production of KmTx-2 in both treatments, but especially in P-limited cultures. Because the KmTx-1 toxin is an order of magnitude more potent than KmTx-2, total cellular toxicity was increased dramatically at high pCO 2 , particularly in P-limited cultures. Specific growth rates were accelerated by enriched CO 2 in P-replete cultures, but not in P-limited treatments. Growth rates or toxicity of K. veneficum could increase substantially in the future with high CO 2 levels in the ocean, depending on P availability, and so interactions between rising CO 2 and eutrophication could cause major shifts in present day patterns of harmful algal toxin production. These results suggest that over the coming decades, rising CO 2 could substantially increase karlotoxin damage to food webs in the often P-limited estuaries where Karlodinium blooms occur.

Laboratory culture experiments were used to investigate the growth rate of colonial Phaeocystis antarctica as a function of irradiance and dissolved iron concentration. The experiments were conducted with a P. antarctica strain isolated from the southern Ross Sea, Antarctica, and made use of natural, low-iron (

Interpretation of photosynthetic pigment data using iterative programs such as CHEMTAX are widely used to examine algal community structure in the surface ocean. The accuracy of such programs relies on understanding the effects of environmental parameters on the pigment composition of taxonomically diverse algal groups. Phaeocystis antarctica is an important contributor to total autotrophic production and the biogeochemical cycling of carbon and sulfur in the Southern Ocean. Here we report the results of a laboratory culture experiment in which we examined the effects of ambient dissolved iron concentration on the pigment composition of colonial P. antarctica, using a new P. antarctica strain isolated from the southern Ross Sea in December 2003. Low-iron ( 3) indicative of Fe stress. We also observed that the ratio of fucoxanthin to 19′-hexanoyloxyfucoxanthin (Fuco:Hex ratio) was highly correlated (r 2 = 0.82) with initial dissolved Fe concentration, with Fuco:Hex ratios

We have carried out field and laboratory experiments to examine the iron requirements of colonial Phaeocystis antarctica in the Ross Sea. In December 2003, we performed an iron/light-manipulation bioassay experiment in the Ross Sea polynya, using an algal assemblage dominated by colonial Phaeocystis antarctica, collected from surface waters with an ambient dissolved Fe concentration of ∼0.4 nM. Results from this experiment suggest that P. antarctica growth rates were enhanced at high irradiance (∼50% of incident surface irradiance) but were unaffected by iron addition, and that elevated irradiance mediated a significant decrease in cellular chlorophyll a content. We also conducted a laboratory iron dose–response bioassay experiment using a unialgal, non-axenic strain of colonial P. antarctica and low-iron (

We examined the physiological responses of steady-state iron (Fe)-replete and Fe-limited cultures of the biogeochemically critical marine unicellular diazotrophic cyanobacterium Crocosphaera at glacial (19 Pa; 190 ppm), current (39 Pa; 380 ppm), and projected year 2100 (76 Pa; 750 ppm) CO 2 levels. Rates of N 2 and CO 2 fixation and growth increased in step with increasing partial pressure of CO 2 (pCO 2 ), but only under Fe-replete conditions. N 2 and carbon fixation rates at 75 Pa CO 2 were 1.4-1.8-fold and 1.2-2.0-fold higher, respectively, relative to those at present day and glacial pCO 2 levels. In Fe-replete cultures, cellular Fe and molybdenum quotas varied threefold and were linearly related to N 2 fixation rates and to external pCO 2 . However, N 2 fixation and trace metal quotas were decoupled from pCO 2 in Fe-limited Crocosphaera. Higher CO 2 and Fe concentrations both resulted in increased cellular pigment contents and affected photosynthesis vs. irradiance parameters. If these results also apply to natural Crocosphaera populations, anthropogenic CO 2 enrichment could substantially increase global oceanic N 2 and CO 2 fixation, but this effect may be tempered by Fe availability. Possible biogeochemical consequences may include elevated inputs of new nitrogen to the ocean and increased potential for Fe and/or phosphorus limitation in the future high-CO 2 ocean, and feedbacks to atmospheric pCO 2 in both the near future and over glacial to interglacial timescales.

Biological N2 fixation is the dominant supply of new nitrogen (N) to the oceans, but is often inhibited in the presence of fixed N sources such as nitrate (NO3 −). Anthropogenic fixed N inputs to the ocean are increasing, but their effect on marine N2 fixation is uncertain. Thus, global estimates of new oceanic N depend on a fundamental understanding of factors that modulate N source preferences by N2-fixing cyanobacteria. We examined the unicellular diazotroph Crocosphaera watsonii (strain WH0003) to determine how the light-limited growth rate influences the inhibitory effects of fixed N on N2 fixation. When growth (µ) was limited by low light (µ = 0.23 d−1), short-term experiments indicated that 0.4 µM NH4 + reduced N2-fixation by ∼90% relative to controls without added NH4 +. In fast-growing, high-light-acclimated cultures (µ = 0.68 d−1), 2.0 µM NH4 + was needed to achieve the same effect. In long-term exposures to NO3 −, inhibition of N2 fixation also varied with growth rate. In high-light-acclimated, fast-growing cultures, NO3 − did not inhibit N2-fixation rates in comparison with cultures growing on N2 alone. Instead NO3 − supported even faster growth, indicating that the cellular assimilation rate of N2 alone (i.e. dinitrogen reduction) could not support the light-specific maximum growth rate of Crocosphaera. When growth was severely light-limited, NO3 − did not support faster growth rates but instead inhibited N2-fixation rates by 55% relative to controls. These data rest on the basic tenet that light energy is the driver of photoautotrophic growth while various nutrient substrates serve as supports. Our findings provide a novel conceptual framework to examine interactions between N source preferences and predict degrees of inhibition of N2 fixation by fixed N sources based on the growth rate as controlled by light.