Redfield Revisited: variability of C:N:P in marine microalgae and its biochemical basis

A compilation of data on the elemental composition of marine phytoplankton from published studies was used to determine the range of C:N:P. The N:P ratio of algae and cyanobacteria is very plastic in nutrient-limited cells, ranging from <5 mol N:mol P when phosphate is available greatly in excess of nitrate or ammonium to <100 mol N:mol P when inorganic N is present greatly in excess of P. Under optimal nutrient-replete growth conditions, the cellular N:P ratio is somewhat more constrained, ranging from 5 to 19 mol N:mol P, with most observations below the Redfield ratio of 16. Limited data indicate that the critical N:P that marks the transition between N- and P-limitation of phytoplankton growth lies in the range 20–50 mol N:mol P, considerably in excess of the Redfield ratio. Biochemical composition can be used to constrain the critical N:P. Although the biochemical data do not preclude the critical N:P from being as high as 50, the typical biochemical composition of nutrient-replete algae and cyanobacteria suggests that the critical N:P is more likely to lie in the range between 15 and 30. Despite the observation that the overall average N:P composition of marine particulate matter closely approximates the Redfield ratio of 16, there are significant local variations with a range from 5 to 34. Consistent with the culture studies, lowest values of N:P are associated with nitrate- and phosphate-replete conditions. The highest values of N:P are observed in oligotrophic waters and are within the range of critical N:P observed in cultures, but are not so high as to necessarily invoke P-limitation. The C:N ratio is also plastic. The average C:N ratios of nutrientreplete phytoplankton cultures, oceanic particulate matter and inorganic N and C draw-down are slightly greater than the Redfield ratio of 6.6. Neither the analysis of laboratory C:N:P data nor a more theoretical approach based on the relative abundance of the major biochemical molecules in the phytoplankton can support the contention that the Redfield N:P reflects a physiological or biochemical constraint on the elemental composition of primary production

Role of zooplankton dynamics for Southern Ocean phytoplankton biomass and global biogeochemical cycles

Global ocean biogeochemistry models currently employed in climate change projections use highly simplified representations of pelagic food webs. These food webs do not necessarily include critical pathways by which ecosystems interact with ocean biogeochemistry and climate. Here we present a global biogeochemical model which incorporates ecosystem dynamics based on the representation of ten plankton functional types (PFTs); six types of phytoplankton, three types of zooplankton, and heterotrophic bacteria. We improved the representation of zooplankton dynamics in our model through (a) the explicit inclusion of large, slow-growing zooplankton, and (b) the introduction of trophic cascades among the three zooplankton types. We use the model to quantitatively assess the relative roles of iron vs. grazing in determining phytoplankton biomass in the Southern Ocean High Nutrient Low Chlorophyll (HNLC) region during summer. When model simulations do not represent crustacean macrozooplankton grazing, they systematically overestimate Southern Ocean chlorophyll biomass during the summer, even when there was no iron deposition from dust. When model simulations included the developments of the zooplankton component, the simulation of phytoplankton biomass improved and the high chlorophyll summer bias in the Southern Ocean HNLC region largely disappeared. Our model results suggest that the observed low phytoplankton biomass in the Southern Ocean during summer is primarily explained by the dynamics of the Southern Ocean zooplankton community rather than iron limitation. This result has implications for the representation of global biogeochemical cycles in models as zooplankton faecal pellets sink rapidly and partly control the carbon export to the intermediate and deep ocean

20 years of the Atlantic Meridional Transect - AMT

The AMT (www.amt-uk.org) is a multidisciplinary programme which undertakes biological, chemical, and physical oceanographic research during an annual voyage between the UK and a destination in the South Atlantic such as the Falkland Islands, South Africa, or Chile. This transect of >12,000 km crosses a range of ecosystems from subpolar to tropical, from euphotic shelf seas and upwelling systems, to oligotrophic mid-ocean gyres. The year 2015 has seen two milestones in the history of the AMT: the achievement of 20 years of this unique ocean going programme and the departure of the 25th cruise on the 15th of September. Both of these events were celebrated in June this year with an open science conference hosted by the Plymouth Marine Laboratory (PML) and will be further documented in a special issue of Progress in Oceanography which is planned for publication in 2016. Since 1995, the 25 research cruises have involved 242 sea-going scientists from 66 institutes representing 22 countries. AMT was designed from the outset to be a collaborative programme. It was originally conceived by Jim Aiken, Patrick Holligan, Roger Harris, and Dave Robins with Chuck McClain and Chuck Trees at NASA to test and ground truth satellite algorithms of ocean color. The opportunities offered by this initiative meant that this series of repeated biannual cruises rapidly developed into a coordinated study of ocean biodiversity, biogeochemistry, and ocean/atmosphere interactions

Control of light saturated photosynthesis: Concentration and activity of ribulose bisphosphate carboxylase. Final report, September 1, 1993--February 28, 1997

Ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) is one of the most abundant enzymes on the planet and is responsible for catalysing the net fixation of CO{sub 2} into organic matter. It is central, therefore, to primary productivity in marine and terrestrial ecosystems. Rubisco is a large enzyme with low substrate affinity and low catalytic efficiency and is considered to limit the rate of light-saturated photosynthesis. This report summarizes research into the molecular basis of the regulation of phytoplankton photosynthesis. It describes experimental and theoretical studies of the role of Rubisco in regulating the photosynthetic rate of phytoplankton. It also describes the integration of a mechanistically based phytoplankton growth model into a description of primary productivity in the sea. This work was conducted as part of the Ocean Margins Program

Response of the photosynthetic apparatus of Phaeodactylum tricornutum (Bacillariophyceae) to nitrate, phosphate or iron starvation.

The effects of nitrate, phosphate, and iron starvation and resupply on photosynthetic pigments, selected photosynthetic proteins, and photosystem II (PSII) photochemistry were examined in the diatom Phaeodactylum tricornutum Bohlin (CCMP 1327). Although cell chlorophyll a (chl a) content decreased in nutrient‐starved cells, the ratios of light‐harvesting accessory pigments (chl c and fucoxanthin) to chl a were unaffected by nutrient starvation. The chl a‐specific light absorpition coefficient (a*) and the functional absorption cross‐section of PSII (σ) increased during nutrient starvation, consistent with reduction of intracellular self‐shading (i.e. a reduction of the “package effect”) as cells became chlorotic. The light‐harvesting complex proteins remained a constant proportion of total cell protein during nutrient starvation, indicating that chlorosis mirrored a general reduction in cell protein content. The ratio of the xanthophylls cycle pigments diatoxanthin and diadinoxanthin to chl a increased during nutrient starvation. These pigments are thought to play a photo‐protective role by increasing dissipation of excitation energy in the pigment bed upstream from the reaction centers. Despite the increase in diatoxanthin and diadinoxanthin, the efficiency of PSII photochemistry, as measured by the ration of variable to maximum fluorescence (Fv/Fm) of dark‐adapted cells, declined markedly under nitrate and iron starvation and moderately under phosphate starvation. Parallel to changes in Fv/Fm were decreases in abundance of the reaction center protein D1 consistent with damage of PSII reaction centers in nutrient‐starved cells. The relative abundance of the carboxylating enzyme, ribulose bisphosphate carboxylase/oxygenase (RUBISCO), decreased in response to nitrate and iron starvation but not phosphate starvation. Most marked was the decline in the abundance of the small subunit of RUBISCO in nitrate‐starved cells. The changes in pigment content and fluorescence characteristics were typically reversed within 24 h of resupply of the limiting nutrient

Induction of specific proteins in eukaryotic algae grown under iron, phosphorus and nitrogen deficient conditions

What limits phytoplankton growth in nature? The answer is elusive because of methodological problems associated with bottle incubations and nutrient addition experiments. We are investigating the possibility that antibodies to proteins repressed by a specific nutrient can be used as probes to indicate which nutrient limits photosynthetic carbon fixation in the ocean. The diatom Phaeodactylum tricornutum Bohlin and the chlorophyte Dunaliella tertiolecta Butcher were grown in batch cultures in artificial seawater and f/2 nutrient lacking either phosphorus, iron, or nitrogen. Chlorosis was induced by nutrient limitation in both species with the exception of phosphorus‐limited D. tertiolecta. The synthesis and appearance of specific proteins were followed by labeling with 14C‐bicarbonate. Nutrient limitation in general leads to a decrease in the quantum efficiency of photosystem II, suggesting that deficiency of any nutrient affects the photosynthetic apparatus to some degree: however, the effect of nitrogen and iron limitation on quantum efficiency is more severe than that of phosphorus. A crude fractionation of the soluble and membrane proteins demonstrated that the large proteins induced under limitation by phosphorus and iron were associated with the membranes. However, small iron‐repressible proteins were located in the soluble fraction. Isolation with anion‐exchange chromatography and N‐terminal sequencing of iron‐repressible, 23‐kDa Proteins from D. tertiolecta, P. tricornutum, and Chaetoceros gracilis revealed that these small soluble proteins have strong homology with the N‐terminal sequence of flavodoxins from Azotobacter and Clostridium. The identity of the flavodoxin from D. tertiolecta was confirmed by immunodetection using antiflavodoxin raised against Chlorella. Flavodoxin was detected only under iron deprivation and was absent from nitrogen‐and phosphorus‐limited algae. Flavodoxin is a prime candidate for a molecular probe of iron limitation in the ocean. The requirements to confirm its utility in nature are discussed

Relative influence of nitrogen and phosphorous availability on phytoplankton physiology and productivity in the oligotrophic sub-tropical North Atlantic Ocean

Nutrient addition bioassay experiments were performed in the low-nutrient, low-chlorophyll oligotrophic subtropical North Atlantic Ocean to investigate the influence of nitrogen (N), phosphorus (P), and/or iron (Fe) on phytoplankton physiology and the limitation of primary productivity or picophytoplankton biomass. Additions of N alone resulted in 1.5–2 fold increases in primary productivity and chlorophyll after 48 h, with larger (~threefold) increases observed for the addition of P in combination with N (NP). Measurements of cellular chlorophyll contents permitted evaluation of the physiological response of the photosynthetic apparatus to N and P additions in three picophytoplankton groups. In both Prochlorococcus and the picoeukaryotes, cellular chlorophyll increased by similar amounts in N and NP treatments relative to all other treatments, suggesting that pigment synthesis was N limited. In contrast, the increase of cellular chlorophyll was greater in NP than in N treatments in Synechococcus, suggestive of NP co-limitation. Relative increases in cellular nucleic acid were also only observed in Synechococcus for NP treatments, indicating co-limitation of net nucleic acid synthesis. A lack of response to relief of nutrient stress for the efficiency of photosystem II photochemistry, Fv :Fm, suggests that the low nutrient supply to this region resulted in a condition of balanced nutrient limited growth, rather than starvation. N thus appears to be the proximal (i.e. direct physiological) limiting nutrient in the oligotrophic sub-tropical North Atlantic. In addition, some major picophytoplankton groups, as well as overall autotrophic community biomass, appears to be co-limited by N and P

Phytoplankton photoacclimation and photoadaptation in response to environmental gradients in a shelf sea

Variability in the photosynthetic performance of natural phytoplankton communities, due to both taxonomic composition and the physiological acclimation of these taxa to environmental conditions, was assessed at contrasting sites within a temperate shelf sea region. Physiological parameters relating to the structure of the photosystem II (PSII) antenna and processes downstream from PSII were evaluated using a combination of fast repetition rate fluorescence, oxygen flash yields, spectral fluorescence, and 14C photosynthesis versus irradiance measurements. Parameters relating to PSII antenna structure, specifically the functional absorption cross-section (sPSII) and the chlorophyll to PSII reaction center ratio, varied principally as a result of spatial (horizontal) taxonomic differences. Phenotypic plasticity in the size of the PSII light-harvesting antenna appeared to be limited. In contrast, parameters related to electron transport rates (ETRs) downstream of PSII, including the maximum ETR (1/tPSII), the chlorophyll-specific maximum rate of carbon fixation (P*max), and the light-saturation intensity (Ek), all decreased from the surface to the subsurface chlorophyll maximum (SCM) in stratified waters. The primary photoacclimation response to the vertical light gradient thus resulted in decreasing light-saturated carbon fixation per reaction center with increasing depth. Increases in the ratio of PSII reaction centers to carbon fixation capacity thus dominated the phenotypic response to decreased irradiance within the SCM. Perhaps counterintuitively, phytoplankton populations within fully mixed water columns, characterized by low mean irradiance, were acclimated or adapted to relatively high irradiance. <br/

Nitrogen and phosphorus co-limitation of bacterial productivity and growth in the oligotrophic subtropical North Atlantic

Bacterial productivity and biomass are thought to be limited by dissolved organic carbon (DOC) in much of the world?s oceans. However, the mixed layer of oligotrophic oceans is often depleted in dissolved inorganic nitrogen and phosphate, raising the possibility that macronutrients may also limit heterotrophic bacterial growth. We used nutrient bioassay experiments to determine whether inorganic nutrients (N, P, Fe) and/or DOC could limit bacterial productivity and biomass in the central North Atlantic during the spring of 2004 (Mar-Apr). We observed that both heterotrophic bacterial productivity and biomass were co-limited by N and P in the oligotrophic North Atlantic, and additions of labile DOC (glucose) provided no stimulation unless N and P were also added. Flow cytometry results indicated that only a small subset of large cells high in nucleic acid content were responsible for the increased productivity in the combined NP amendments. In contrast, nutrient additions elicited no net change on the dominant component of the bacterial population, composed of small cells with relatively low nucleic acid content. In the combined NP treatments the relative increase in bacterial production was greater than that measured when phytoplankton productivity was relieved of nitrogen limitation. These results suggest that N and P co-limitation in the bacterial community results in increased competition between the heterotrophic and autotrophic components of the surface communities in the Central North Atlantic Ocean, and potentially impacts the cycling of organic matter by the bacterioplankton