Marine diatoms grown in chemostats under silicate or ammonium limitation. III. Cellular chemical composition and morphology of Chaetoceros debilis,Skeletonema costatum,and Thalassiosira gravida

Three marine diatoms, Skeletonema costatum, Chaetoceros debilis, and Thalassiosira gravida were grown under no limitation and ammonium or silicate limitation or starvation. Changes in cell morphology were documented with photomicrographs of ammonium and silicate-limited and non-limited cells, and correlated with observed changes in chemical composition. Cultures grown under silicate starvation or limitation showed an increase in particulate carbon, nitrogen and phosporus and chlorophyll a per unit cell volume compared to non-limited cells; particulate silica per cell volume decreased. Si-starved cells were different from Si-limited cells in that the former contained more particulate carbon and silica per cell volume. The most sensitive indicator of silicate limitation or starvation was the ratio C:Si, being 3 to 5 times higher than the values for non-limited cells. The ratios Si:chlorophyll a and S:P were lower and N:Si was higher than non-limited cells by a factor of 2 to 3. The other ratios, C:N, C:P, C:chlorophyll a, N:chlorophyll a, P:chlorophyll a and N:P were considered not to be sensitive indicators of silicate limitation or starvation. Chlorophyll a, and particulate nitrogen per unit cell volume decreased under ammonium limitation and starvation. NH₄-starved cells contained more chlorophyll a, carbon, nitrogen, silica, and phosphorus per cell volume than NH₄-limited cells. N:Si was the most sensitive ratio to ammonium limitation or starvation, being 2 to 3 times lower than non-limited cells. Si:chlorophyll a, P:chlorophyll a and N:P were less sensitive, while the ratios C:N, C:chlorophyll a, N:chlorophyll a, C:Si, C:P and Si:P were the least sensitive. Limited cells had less of the limiting nutrient per unit cell volume than starved cells and more of the non-limiting nutrients (i.e., silica and phosphorus for NH₄-limited cells). This suggests that nutrient-limited cells rather than nutrient-starved cells should be used along with non-limited cells to measure the full range of potential change in cellular chemical composition for one species under nutrient limitation.Contribution No. 943 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Resting spore formation and biochemical composition of the marine planktonic diatom Chaetoceros pseudocurvisetus in culture: ecological significance of decreased nucleotide content and activation of the xanthophyll cycle by resting spore formation

The biochemical composition of vegetative cells and resting spores of Chaetoceros pseudocurvisetus Mangin was compared in cultures under various nutrient and light conditions. The cellular content of major nucleotides such as AMP, ADP, ATP and UTP decreased in the order: vegetative cells, nutrient-starved (vegetative) cells and resting spores, indicating that the general metabolism of resting spores is relatively inactive. ADP-glucose was only abundant in nutrient-starved vegetative cells, suggesting metabolic imbalance in these cells. The chl a–specific fluorescence yield of vegetative cells grown under all culture conditions was low, but very high in resting spores. The ratios of the cellular contents of diadinoxanthin to chl a and of diatoxanthin to chl a were higher in resting spores and nutrient-starved vegetative cells than in nutrient-replete vegetative cells. The diadinoxanthin–diatoxanthin xanthophyll cycle was active in resting spores; the xanthophyll cycle was synchronized with a 14 h light:10 h dark photoperiod. Also, the ratios of cellular content of diadinoxanthin and diatoxanthin to cellular content of chl a in resting spores were relatively high in high irradiance, and decreased gradually in conditions of darkness over long culture periods. Under conditions of strong light and high temperature, most resting spores survived more than 40 d while nutrient-starved vegetative cells died within 33 d. These results suggest that resting spore formation is a strategy for enhancing protection and lowering metabolic rate for survival. These physiological changes accompanying spore formation enable resting spores not only to overwinter but also to “oversummer” in the coastal euphotic layer. Received: 23 March 1999 / Accepted: 11 August 1999     

Dynamique du phytoplancton et matière organique particulaire dans la zone euphotique de l'Atlantique Equatorial

At two fixed stations in the Equatorial Atlantic Ocean (0°–4° W), the physical, chemical and biological properties of the euphotic layer were determined for 14 d (Station A: 5–18 February, 1979) and 13 d (Station B: 20 October–7 November, 1979), respectively. The stability of the water column allowed comparison of 3 different “systems”: (i) a well-illuminated and nitrate-depleted mixed layer; (ii) a chlorophyll maximum layer (chl a _max) in the thermocline which is poorly illuminated (6.3% of surface irradiance); (iii) a well-illuminated but nitrate-rich (>0.9 μg-at l^-1) mixed layer. In each layer the particulate organic carbon (COP), nitrogen (NOP) and phosphorus (POP) contents were measured and compared with the phytoplankton biomass. In the chlorophyll maximum layer, the phytoplankton biomass contributed significantly to the total particulate organic matter (between 55 and 75%). In the nitrate-depleted mixed layer, the results varied according to whether the regression technique [COP=f(chl a)] was used, or the chl a synthesis during the incubation of the samples. With the former technique, the phytoplankton carbon (C _p) content appeared minimal, because the y intercept, computed using all the data of the water column, was probably overestimated for this layer. POP would be more associated with living protoplasm than with carbon and nitrogen in the three layers. In the chlorophyll a maximum layer it constitutes a valuable detritus-free biomass measurement, since 80% of the POP consist of phytoplankton phosphorus. The assimilation numbers (NA=μg C μg chl a ^-1 h^-1) were high in all three layers, but the highest values were recorded in the nitrate-depleted mixed layer (NA=15 μg C μg chl a ^-1 h^-1). In the chlorophyll maximum layer, light would be a limiting factor during incubation: between 10²⁵ and 8.10²⁴ quanta m^-2 d^-1 NA and light are positively correlated independant of nitrate concentration. The growth rates of phytoplankton (μ) were estimated and compared to the maximum expected growth rate. Our main conclusion was that despite very low biomass and nutrient content, the mixed layer was in a highly dynamic state, as evidenced by high rates of phytoplankton growth and short nutrient turnover times (1 d or less for PO-P₄ in the mixed layer versus 3 d in the thermocline). The presence of nitrate in the water column allows the development of a higher phytoplankton biomass but does not increase growth rate.     

A new method for estimating phytoplankton growth rates and carbon biomass

A new method is described for the determination of phytoplankton growth rates and carbon biomass. This procedure is easy to apply and utilizes the labeling of chlorophyll a (chl a) with ¹⁴C. Pure chl a is isolated using two-way thin-layer chromatography, and the specific activity of chl a carbon is determined. Data from laboratory cultures indicate that the specific activity of chl a carbon becomes nearly equal to that of total phytoplankton carbon in incubations lasting 6 to 12 h and can be used to calculate phytoplankton growth rates and carbon biomass. Application of the method to the phytoplankton community in an eutrophic estuary in Hawaii indicates that the cells are growing with a doubling time of about 2 d and that about 85% of the particulate carbon consists of phytoplankton carbon.     

Survival and recovery of resting spores and resting cells of the marine planktonic diatom Chaetoceros pseudocurvisetus under fluctuating nitrate conditions

Of the two resting life-forms of the planktonic diatom Chaetoceros pseudocurvisetus Mangin formed during periods of nitrate depletion, resting spores survived at least 1 month after spore formation at 24 °C, while resting cells survived only for about 10 d at the same temperature. Under nitrogen limitation, resting cells exhibited higher specific death rates than resting spores at temperatures ranging from 5 to 30 °C. After nitrogen replenishment, resting spores required a certain lag period of about 1 d to initiate vegetative growth at levels of nitrate supply from 0.5 to 20 M, while resting cells initiated vegetative growth almost immediately. Resting spores exhibited an intracellular accumulation of the supplied nitrate during germination and initial vegetative growth. The resting cells, however, exhibited more active vegetative growth, closely coupled with the uptake of the supplied nitrate. The resting spores and resting cells appear to play different roles in the maintenance of populations under nutrient fluctuations depending on the interval length between nutrient fluxes in natural waters. Received: 27 April 1998 / Accepted: 1 March 1999     

Nutrient limitation in the giant clam-zooxanthellae symbiosis: effects of nutrient supplements on growth of the symbiotic partners

The effect of ammonium (5, 10 M N) and phosphate (2, 5, 10 M P) on the growth of the giant clam Tridacna gigas and its symbiotic dinoflagellate Symbiodinium sp. was examined. A 3 mo exposure to these nutritients significantly increased the N or P composition of the soft tissues, as reflected in a corresponding change in C:N:P ratio. Furthermore, exposure to N or N+P markedly increased the amount of soft tissue, but P alone did not, demonstrating that increased availability of inorganic nitrogen enhances tissue growth of the clam host. With addition of N, or N+P, there was a significant increase in the total number of zooxanthellae per clam, with a corresponding decrease in chlorophyll a (chl a) content per zooxanthella. However, only with N+P was there an increase in the zooxanthellae mitotic index. The inverse relationship between zooxanthellae number and chl a per zooxanthella is consistent with phytoplankton studies indicating conditions of nutrient-limitation. Furthermore, the unaffected C:N:P composition of the zooxanthellae and their relatively low specific-growth rates (4 to 10%) also suggest that they are nutrient-limited in vivo. In particular, their high mean C:N:P ratio of 303:52:1 indicates that, relative to C, they are much more depleted in P and less in N than are free-living phytoplankton. Furthermore, polyphosphates (phosphate reserves) were undetectable, and the activity levels of acid phosphatase in the zooxanthellae were relatively high and not influenced by the host's exposure to increased P concentrations in the sea water, implicating the clam host in active regulation of P availability to its symbiotic algae. This is strong evidence that N-limitation of clam zooxanthellae is a function of the availability of ammonium to the symbiosis while, irrespective of nutrient levels in sea water, clam zooxanthellae still show characteristics of P-limitation.     

Floristic and physiological differences between the shallow and the deep nanophytoplankton community in the euphotic zone of the open tropical Atlantic revealed by HPLC analysis of pigments

Suspended matter sampled in 1982 in the North Equatorial Current, in the open Atlantic to the west of West Africa, was analyzed by high performance liquid chromatography. The pigment fingerprint of samples taken in the surface mixed layer was dominated by zeaxanthin and chlorophyll a, in agreement with observed dominance of coccoid cyanobacteria. Near the bottom of the euphotic zone the fingerprint was more complicated, with a sharp transition at the depth of the deep chlorophyll maximum layer to dominance of chlorophyll b, 19-hexanoyloxyfucoxanthin and an unknown fucoxanthin derivative in the lower part of this layer; this fingerprint suggests dominance of eukaryotes (green algae, Prymnesiophyceae and Chrysophyceae) at depth. Up to 90% of the chl a was contained in particles smaller than 8 m, and in the surface mixed layer even more than 50% in particles smaller than 1 m. The high concentration of zeaxanthin relative to chl a near the surface suggests adaptation of the cyanobacteria to exposure to high irradiance. Evidence of this adaptation was the very high specific phytoplankton growth rate between sunrise and sunset (=0.16 h^-1), measured by recording ¹⁴C incorporation into organic carbon and into chl a carbon after isolation of the latter by HPLC. The high concentration of chl b relative to chl a at depth was possibly caused by shade-adapted green algae containing more chl b than chl a. The specific growth rate of the deep shade community was low (<0.04 h^-1), yet net primary production, calculated on the basis of chl a increase during incubation, was greatest at depth.     

Effect of nutrient enrichment in the field on the biomass, growth and calcification of the giant clam Tridacna maxima

Nutrients were added separately and combined to an initial concentration of 10 μM (ammonium) and/or 2 μM (phosphate) in a series of experiments carried out with the giant clam Tridacna maxima at 12 microatolls in One Tree Island lagoon, Great Barrier Reef, Australia (ENCORE Project). These nutrient concentrations remained for 2 to 3 h before returning to natural levels. The additions were made every low tide (twice per day) over 13 and 12 mo periods for the first and second phase of the experiment, respectively. The nutrients did not change the wet tissue weight of the clams, host C:N ratio, protein content of the mantle, calcification rates or growth rates. However, ammonium (N) enrichment alone significantly increased the total population density of the algal symbiont (Symbiodinium sp.: C = 3.6 · 10⁸ cell clam⁻¹, N = 6.6 · 10⁸ cell clam⁻¹, P = 5.7 · 10⁸ cell clam⁻¹, N + P = 5.7 · 10⁸ cell clam⁻¹; and C = 4.1 · 10⁸ cell clam⁻¹, N = 5.1 · 10⁸ cell clam⁻¹, P = 4.7 · 10⁸ cell clam⁻¹, N + P = 4.5 · 10⁸ cell clam⁻¹, at the end of the first and second phases of the experiment, respectively), although no differences in the mitotic index of these populations were detected. The total chlorophyll a (chl a) content per clam but not chlorophyll a per cell also increased with ammonium addition (C = 7.0 mg chl a clam⁻¹, N = 13.1 mg chl a clam⁻¹, P = 12.9 mg chl a clam⁻¹, N + P = 11.8 mg chl a clam⁻¹; and C = 8.8 mg chl a clam⁻¹, N = 12.8 mg chl a clam⁻¹; P = 11.2 mg chl a clam⁻¹, N + P = 11.3 mg chl a clam⁻¹, at the end of the first and second phases of the experiment, respectively). The response of clams to nutrient enrichment was quantitatively small, but indicated that small changes in inorganic nutrient levels affect the clam–zooxanthellae association. Received: 2 June 1997 / Accepted: 9 June 1997     

Resting spore formation and phosphorus composition of the marine diatom Chaetoceros pseudocurvisetus under various nutrient conditions

The marine diatom Chaetoceros pseudocurvisetus was used for a study of resting spore formation and cellular phosphorus composition. Resting spores were found in any culture medium with ample silica, including nitrogen limited, phosphorus limited and nutrient replete conditions. Resting spores protected themselves with thick silica walls, so that vegetative cells required about 3 pmol cell^-1 of additional silica to form resting spores. Phosphorus compounds in the cells were divided into eight fractions: nucleotide-P, orthophosphate, acid soluble polyphosphate, sugar phosphate, nucleic acid-P, acid insoluble polyphosphate, lipid-P and residual-P. The sum of orthophosphate, sugar phosphate and nucleic acid-P comprised over 65% of the total phosphorus in cells under any culture conditions. Sugar phosphate was the most variable component, being most abundant in vegetative cells and least abundant in resting spores.     

Acclimation and adaptation to irradiance in symbiotic dinoflagellates. II. Response of chlorophyll–protein complexes to different photon-flux densities

The response of chlorophyll–protein complexes to super- and sub-saturating photon-flux densities, PFD (250?μmol quanta m^?2?s^?1 and 40?μmol quanta m^?2?s^?1, respectively) were analyzed for Symbiodinium microadriaticum Freudenthal, the symbiont of the Caribbean jellyfish Cassiopeia xamachana; S. kawagutii Trench and Blank, the symbiont of the Indo-Pacific scleractinian Montipora verrucosa; and S. pilosum Trench and Blank, the symbiont of the Carribbean zoanthid Zoanthus sociatus. The results indicate that each species exhibits a quantitatively distinct chlorophyll (chl) a distribution among its chl–protein complexes when cultured under standardized high and low light conditions. In response to sub-saturating PFD, the three species differentially increased the cellular concentrations of most of the chl–protein complexes. Increases in P₇₀₀ (reaction center of Photosystem I) under sub-saturating PFD correlate with an increase in the cellular concentrations of the Photosystem I-enriched complexes. Similarly, increases in photosynthetic unit (PSU) size correlate with an increase in the cellular concentrations of the water-soluble peridinin–chl-a–protein (PCP) complexes and the membrane-bound chl a–chl c ₂–peridinin–protein (acpPC) complexes, which together represent the light-harvesting components of this group. In S. microadriaticum, acclimation to sub-saturating PFD uniquely includes the preferential enrichment of the dimeric form of PCP. Under super-saturating PFD, an enrichment in photo-protective xanthophylls was detected in acpPC from S. microadriaticum and S. pilosum, but not from S. kawagutii. Each species demonstrated a characteristic photo-acclimatory response which correlates with its distribution as endosymbiont in nature, supporting the concept that different species of symbiotic dinoflagellates are adapted (sensu Björkman 1981) to different photic environments. The study was conducted between May 1992 and November 1994.     

Resting spore formation of the marine planktonic diatom Chaetoceros anastomosans induced by high salinity and nitrogen depletion

The marine planktonic diatom Chaetoceros anastomosans, which was isolated from Sagami Bay, was used for a study of resting spore formation mechanisms in batch culture experiments. Vegetative cells could grow at salinities ranging from 20.7 to 45.5‰, and resting spore formation was enhanced significantly in nitrate-depleted, high salinity media (40.0 to 45.5‰). The rate of resting spore formation (1.9 d⁻¹) was comparable to the specific growth rate (1.8 d⁻¹) of vegetative cells in the exponential growth phase in normal salinity medium. The size of resting spores formed under high salinity conditions was smaller than that of spores formed in normal salinity media. Unlike vegetative cells, resting spores seemed to possess some mechanisms to survive over a wider range of salinities by resisting bacterial attacks on their cell walls. Received: 4 August 1996 / Accepted: 27 August 1996     

Carbon-nitrogen relations at whole-cell and free-amino-acid levels during batch growth of Isochrysis galbana (Prymnesiophyceae) under conditions of alternating light and dark

Isochrysis galbana Parke, Strain CCAP 927/1, was grown in ammonium-limited batch culture under a 12 h light: 12 h dark illumination cycle. Samples were taken every 12 h over the 26 d period from lag phase through exponential into stationary phase (no net carbon fixation), with more frequent sampling at points of interest. Exponential cell-specific growth rate was 0.3 to 0.4d^-1. Cell division occurred during the dark phase, while cell volume increase, ammonium uptake, and pigment synthesis occurred during the light. Stationary phase cells were small, and the lag phase was long (5 d) even though the C:N ratio had returned from 18 to 6.5 within 2 d, followed by synthesis of chlorophyll a. Net chlorophyll synthesis ceased within 4 d of exhaustion of the nitrogen source. The chlorophyll c: chlorophyll a ratio remained constant during increasing nitrogen deprivation. Biovolume and carotenoids correlated with carbon biomass. Levels of chlorophyll a correlated poorly with carbon fixation and carbon biomass once the nitrogen source had been exhausted. Except after the addition of ammonium to nitrogen-deprived cells (refeeding), the content of intracellular glutamine and the glutamine: glutamate ratio were low during the dark phase, rising to a plateau within the first 1 h of illumination. Refeeding of cells which had only just exhausted the extracellular nitrogen source resulted in a much smaller increase in glutamine than refeeding of nitrogen-starved (stationary-phase) cells. Nitrogen biomass correlated with the presence of an unidentified intracellular amine.     

Life-form population responses of a marine planktonic diatom,Chaetoceros pseudocurvisetus,to oligotrophication in regionally upwelled water

Life-form population responses of a centric planktonic diatom,Chaetoceros pseudocurvisetus Mangin, were investigated in summer 1986 and 1988 in the Izu Islands, Japan, in regionally upwelled water where nutrient concentration changed from favorable to unfavorable conditions for active growth and reproduction (oligotrophication). Two types of life form were observed: vegetative cells of healthy and unhealthy looking conditions and resting spores. The observed life-form responses were experimentally evaluated along with a depletion of limiting nutrients. The algal population ceased vegetative growth and initiated resting spore formation with a disappearance of limiting nitrate, and this life-form response seemed to be triggered by the decrease of cellular nitrogen content below a certain level. Since a large amount of silicon was required for the resting spore formation, a part of vegetative cells were unable to form resting spores and formed unhealthy looking vegetative cells under insufficient concentrations of silicic acid. Percentage shares of the resting spores in the population were linearly related to the amounts of available silicic acid. Vegetative cells which did not form resting spores showed greater mortality than resting spores by attack of bacteria and protozoa; however, vegetative cells could respond quickly to possible nutrient replenishment.     

Effets de l'eutrophie et de la dessalure sur les populations phytoplanctoniques

A study undertaken in the surface waters of the Gulf of Fos (France), revealed very low salinities, high nutrient concentrations, and very rich phytoplanktonic populations (maximum 7.5·10⁵ cells/l). Very often the phytoplanktonic populations attain abundances 2 to 15 times higher than maximum population densities found in the Mediterranean Sea. Such high cellular concentrations seem to suppress more or less the rates of relative photosynthesis if the concentration of chlorophyll a per 1 million cells is considered. Furthermore, the analysis of particular carbon yields high values. This phenomenon can be explained by the fact that, in addition to organic carbon, other organic forms of carbon were measured, especially some carbonates which are abundant in fresh waters. The C/N ratios found seem to support this explanation.     

The effects of ammonium and phosphate enrichments on clorophyll a,pigment ratio and species composition of phytoplankton of Vineyard Sound

Seawater containing natural phytoplankton populations from Vineyard Sound, USA was enriched in the laboratory with three levels each of ammonium and phosphate and with a combination of ammonium and phosphate which provided three different N:P ratios. The addition of ammonium produced more cells and chlorophyll a than the control or the phosphate enrichments. However, enrichment with ammonium and phosphate, regardless of the N:P ratio, yielded the most cells and chlorophyll a. Thus, nitrogen seems to be the primary limiting nutrient, with phosphate showing secondary limiting effects. The ratios of photosynthetic pigments decreased with the increased chlorophyll a production in the enriched cultures. There were no significant changes in the species composition within the cultures, so that the observed changes in pigment ratio and chlorophyll a content were due to physiological responses.     

Light-shade adaptation by the oceanic dinoflagellates Pyrocystis noctiluca and P. fusiformis

Species-specific rates of photosynthetic carbon uptake (P), chlorophyll a content and P versus irradiance (P-I), have been measured for cells of Pyrocystis noctiluca and P. fusiformis isolated from natural populations collected in the euphotic zone within and below the surface mixed layer in the Sargasso Sea. These same measurements and the assay for ribulose bis-phosphate carboxylase (RuBP-Case), have been made for cultures of P. noctiluca in a 12 h L: 12 h D photoperiod at 9 different constant or at changing light intensities. In nature chl a cell^-1 was constant throughout the euphotic zone. The photosynthetic capacity (P_max), of cells captured below the surface mixed layer was lower by a factor of 10 compared with cells collected from the surface mixed layer. The P_max for P. noctiluca collected and incubated within the surface mixed layer was the same as for cell cultures grown under high light, non nutrient-limiting conditions, suggesting that photosynthesis in the natural system was not nutrient limited. In laboratory cultures under constant low light intensities, chl a cell^-1 increased by a factor of 5 while both P_max and RuBPCase activity decreased by a factor of ca 4 compared with high light intensities. In changing light intensities both P_max and RuBPCase activities were decreased by factors of 4 during low light intervals while chl a cell^-1 approached a constant intermediate value. The change in chl a cell^-1 in response to prolonged exposure to constant low light intensities was first order with a rate constant of 0.33 d^-1. For all irradiance conditions in culture, the P-I dependence could be described by the simple Michaelis-Menten formula. The ratio of P_max to K_I, (the light intensity where P=P_max/2) was a constant with a Coefficient of Variation of 12%: The constancy of this ratio, the parallel changes in RuBPCase activity with P_max and the constant chl a cell^-1 in the Sargasso Sea imply that for P. noctiluca and presumably P. fusiformis in nature, a dark enzymatic step rather than changes in photosynthetic pigment concentrations may regulate the photosynthetic capacity in the changing photic environment.Contribution no. 1141 from McCollum-Pratt Institute and Department of Biology, The Johns Hopkins University. Supported by DOE contract no. EY 76S20 3278, NSF no. OCE 76-02571 and ONR no. N300014-81-C-0062     

Diel periodicity in cellular chlorophyll content in marine diatoms

Intracellular chlorophyll a content is one of the many measurable parameters which displays a diel rhythm in marine phytoplankton. In asynchronous laboratory cultures of the diatom Skeletonema costatum, cellular chlorophylls a and c exhibit periodicities which closely follow the light-dark cycle and are not the result of cell division. The laboratory cultures also exhibit diel rhythms in cellular flourescence properties and carbon: chlorophyll a ratios. The occurrence of similar patterns of cellular flourescence, carbon: chlorophyll a ratios, and in situ flourescence in diatom-dominated natural phytoplankton communities suggests the possibility of diel rhythms in cellular chlorophyl a content in diatoms in the sea. The data also suggest that the observed periodicity in cellular chlorophyll content is regulated by the diel light cycle and that the co-occurrence of synchronous or phased cell division would only modify the observed periodicity.This research was performed under the auspices of the United States Department of Energy under Contract No. EY-76-C-02-0016     

Flux capacities and acclimation costs in Trichodesmium from the Gulf of Mexico

Phytoplankton function and acclimation are driven by catalytic protein complexes that mediate key physiological transformations, including generation of photosynthetic ATP and reductant, and carbon and nitrogen fixation. Quantitation of capacities for these processes allows estimation of rates for key ecosystem processes, and identification of factors limiting primary productivity. We herein present molar quantitations of PSI, PSII, ATP synthase, RuBisCO and the Fe protein of nitrogenase of Trichodesmium collected from the Gulf of Mexico, in comparison to determinations for a range of cyanobacteria growing in culture. Using these measurements, estimates were generated for Trichodesmium capacities for carbon fixation of 1–3.4 g C g chl a ⁻¹ h⁻¹ and nitrogen fixation of 0.06–0.17 g N g chl a ⁻¹ h⁻¹, with diel variations in capacities. ATP synthase levels show that ATP synthesis capacity is sufficient to support these levels of carbon and nitrogen fixation, and that ATP synthase levels change over the day in accordance with the ATP demands of nitrogenase and RuBisCO activity. Levels of measured complexes indicate that Trichodesmium manifests n-type diel light acclimation through rapid changes in RuBisCO:PSII, supported by significant investment of cellular nitrogen. The plasticity in the levels and stoichiometry of these core complexes show that changes in the abundance of core protein complexes are an important component of acclimation and regulation of metabolic function by Trichodesmium populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Light absorption,photosynthesis and growth of Nannochloris atomus in nutrient-saturated cultures

Nannochloris atomus was maintained in exponential growth at photon flux densities (PFD) from 400 to 700 nm, ranging from 10 to 200 mol m^-2 s^-1. Growth was lightsaturated at PFDs in excess of 100 mol m^-2 s^-1, with a mean light-saturated growth rate at 23 °C of 1.5×10^-5s^-1 (1.2 d^-1). The light-limited growth rates extrapolated to a compensation PFD for growth that was not significantly different from zero, although no changes in cell numbers were observed in a single culture incubated at a PFD of 1.0 mol m^-2s^-1. Dark-respiration rates were independent of PFD, averaging 1.7×10^-6 mol O₂ mol^-1 C s^-1 (0.14 mol O₂ mol^-1 C d^-1). The maximum photon (quantum) efficiency of photosynthesis was also independent of PFD, with a mean value of 0.12 mol O₂ mol^-1 photon. The chlorophyll a-specific light absorption cross-section ranged from 3 to 6×10^-3 m² mg^-1 chl a and was lowest at low PFDs due to intracellular self-shading of pigments associated with high cell-chlorophyll a contents. The C:chl a ratio increased from 10 to 40 mg C mg^-1 chl a between PFDs of 14 and 200 mol m^-2 s^-1. These new observations for N. atomus are compared with our previous observations for the diatom Phaeodactylum tricornutum in terms of an energy budget for microalgal growth.     

Aspects of the biology of resting spores of Thalassiosira nordenskioeldii and Detonula confervacea

Large numbers of resting spores of Thalassiosira nordenskioeldii Cleve and Detonula confervacea (Cleve) Gran were produced when these species were grown at low levels of ammonia and nitrate. The production of resting spores by T. nordenskioeldii was inversely related to temperature. At 0° and 5°C between 68 and 96% of the total cells were resting spores, while at 15°C resting spores were not produced. Resting spores of both species would not survive 7 days in the dark at 20°C. At 0°, 5°, 10° and 15°C, the length of time that the resting spores of both T. nordenskioeldii and D. confervacea remained viable was inversely related to temperature. At 0°C, T. nordenskioeldii remained viable for 576 days. The data suggest that the production of resting spores by these two species does not aid them in the survival of unfavorably high temperatures such as are found in temperate estuaries during the summer. Rather, they appear to be an adaptation for the survival of long periods of darkness in polar regions.