GROWTH AND PHOTOSYNTHESIS OF MARINE SYNECHOCOCCUS (CYANOPHYCEAE) UNDER IRON STRESS

Prokaryotic picoplankton such as Synechococcus are relatively abundant in putatively Fe-limited high-nutrient, low-chlorophyll (HNLC) regions of the oceans. The physiology of Synechococcus under Fe stress has been studied less than eukaryotic algae. Recent evidence suggests that although biomass and growth rates of Synechococcus are not typically Fe limited in situ, cells may still exhibit symptoms of Fe stress. We grew Synechococcus A2169 and WH7803 in laboratory batch cultures in the artificial medium Aquil and enriched natural seawater, at a series of Fe concentrations and Fe:macronutrient ratios, and with either nitrate or ammonium as the sole nitrogen source. Cell yields, and in some experiments exponential specific growth rate (μ), were more readily Fe limited in the Atlantic isolate WH7803 than in the equatorial Pacific isolate A2169. In both strains, final cell yields spanned about an order of magnitude and decreased continuously with Fe concentration from 900 to 3.6 nM (150 μM N, 10 μM P), whereas μ decreased much less and only at Fe concentrations below 90 nM. Synechococcus yield was controlled by both absolute Fe concentration and Fe:macronutrient ratio, but μ was determined primarily by absolute Fe concentration. Contrary to theoretical predictions, neither yield nor μ was higher in Fe-limited cells grown in ammonium compared to nitrate. Under severe Fe stress, cellular chlorophyll (Chl) content and light-saturated gross photosynthetic capacity (P^cell_m) decreased proportionately, and dark respiration (R^cell_d) increased, such that net P^cell_m was extremely low but gross P^Chl_m was unchanged. This is the first report of an absolute increase in R^cell_d under Fe stress in phytoplankton.     

GROWTH REGULATION OF rRNA CONTENT IN PROCHLOROCOCCUS AND SYNECHOCOCCUS (MARINE CYANOBACTERIA) MEASURED BY WHOLE‐CELL HYBRIDIZATION OF rRNA‐TARGETED PEPTIDE NUCLEIC ACIDS1

The cyanobacteria Synechococcus and Prochlorococcus are important primary producers in marine ecosystems. Because currently available approaches for estimating microbial growth rates can be difficult to apply in the field, we have been exploring the feasibility of using quantitative rRNA measurements as the basis for making such estimates. In this study we examined the relationship between rRNA and growth rate in several Synechococcus and Prochlorococcus strains over a range of light‐regulated growth rates. Whole‐cell hybridization with fluorescently labeled peptide nucleic acid (PNA) probes was used in conjunction with flow cytometry to quantify rRNA on a per cell basis. This PNA probing technique allowed rRNA analysis in a phycoerythrin‐containing Synechococcus strain (WH7803) and in a non–phycoerythrin‐containing strain and in Prochlorococcus. All the strains showed a qualitatively similar tri‐phasic relationship between rRNA·cell^?1 and growth rate, involving relatively little change in rRNA·cell^?1 at low growth rates, linear increase at intermediate growth rates, and a plateau and/or decrease at the highest growth rates. The onset of each phase was associated with the relative, rather than absolute, growth rate of each strain. In the Synechococcus strains, rRNA normalized to flow cytometrically measured forward angle light scatter (an indicator of size) was well‐correlated with growth rate across strains. These findings support the idea that cellular rRNA may be useful as an indicator of in situ growth rate in natural Synechococcus and Prochlorococcus populations.     

RELATIONSHIP BETWEEN CELL CYCLE AND LIGHT‐LIMITED GROWTH RATE IN OCEANIC PROCHLOROCOCCUS (MIT9312) AND SYNECHOCOCCUS (WH8103) (CYANOBACTERIA)1

The relationships between growth rate, cell‐cycle parameters, and cell size were examined in two unicellular cyanobacteria representative of open‐ocean environments: Prochlorococcus (strain MIT9312) and Synechococcus (strain WH8103). Chromosome replication time, C, was constrained to a fairly narrow range of values (～4–6 h) in both species and did not appear to vary with growth rate. In contrast, the pre‐ and post‐DNA replication periods, B and D, respectively, decreased with increasing growth rate from maxima of ～30 and 10–20 h to minima of ～4–6 and 2–3 h, respectively. The combined duration of the chromosome replication and postreplication periods (C+D), a quantity often used in the estimation of Prochlorococcus in situ growth rates, varied ～2.4‐fold over the range of growth rates examined. This finding suggests that assumptions of invariant C+D may adversely influence Prochlorococcus growth rate estimates. In both strains, cell mass was the greatest in slowly growing cells and decreased 2‐ to 3‐fold over the range of growth rates examined here. Estimated cell mass at the start of replication appeared to decrease with increasing growth rate, indicating that the initiation of chromosome replication in Prochlorococcus and Synechococcus is not a simple function of cell biomass, as suggested previously. Taken together, our results reflect a notable degree of similarity between oceanic Synechococcus and Prochlorococcus strains with respect to their growth‐rate‐specific cell‐cycle characteristics.     

INTERACTIVE EFFECTS OF IRRADIANCE AND CO2 ON CO2 FIXATION AND N2 FIXATION IN THE DIAZOTROPH TRICHODESMIUM ERYTHRAEUM (CYANOBACTERIA)1

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 < 0.05) in response to high pCO₂ under mid‐ and high irradiances only. These data imply that the stimulatory effect of elevated pCO₂ on CO₂ fixation and N₂ fixation by T. erythraeum is correlated with light. The ratio of gross:net N₂ fixation was also correlated with light and trichome length in IMS101. Our study suggests that elevated pCO₂ may have a strong positive effect on Trichodesmium gross N₂ fixation in intermediate and bottom layers of the euphotic zone, but perhaps not in light‐saturated surface layers. Climate change models must consider the interactive effects of multiple environmental variables on phytoplankton and the biogeochemical cycles they mediate.     

COMBINED EFFECTS OF TEMPERATURE AND IRON ON THE GROWTH AND PHYSIOLOGY OF THE MARINE DIATOM PHAEODACTYLUM TRICORNUTUM (BACILLARIOPHYCEAE)

Phaeodactylum tricornutum Bohlin (Bacillariophyceae) was maintained in exponential growth under Fe‐replete and stressed conditions over a range of temperatures from 5 to 30° C. The maximum growth rate (GR) was observed at 20° C (optimal temperature) for Fe‐replete and ‐stressed cells. There was a gradual decrease in the GR decreasing temperatures below the optimum temperature; however, the growth rate dropped sharply as temperature increased above the optimum temperature. Fe‐stressed cells grew at half the growth rate of Fe‐replete cells at 20° C, whereas this difference became larger at lower temperatures. The change in metabolic activities showed a similar pattern to the change in growth rate temperature aside from their optimum temperature. Nitrate reductase activity (NRA) and respiratory electron transport system activity (ETS) per cell were maximal between 15 and 20° C, whereas cell‐specific photosynthetic rate (P_cell) was maximal at 20° C for Fe‐replete cells. These metabolic activities were influenced by Fe deficiency, which is consistent with the theoretical prediction that these activities should have an Fe dependency. The degree of influence of Fe deficiency, however, was different for the four metabolic activities studied: NRA > P_cell > ETS = GR. NRA in Fe‐stressed cells was only 10% of that in Fe‐replete cells at the same temperature. These results suggest that cells would have different Fe requirements for each metabolic pathway or that the priority of Fe supply to each metabolic reaction is related to Fe nutrition. In contrast, the order of influence of decreasing the temperature from the optimum temperature was ETS > P_cell > NRA > GR. For NRA, the observed temperature dependency could not be accounted for by the temperature dependency of the enzyme reaction rate itself that was almost constant with temperature, suggesting that production of the enzyme would be temperature dependent. For ETS, both the enzyme reactivity and the amount of enzyme accounted for the dependency. This is the first report to demonstrate the combined effects of Fe and temperature on three important metabolic activities (NRA, P_cell, and ETS) and to determine which activity is affected the most by a shortage of Fe. Cellular composition was also influenced by Fe deficiency, showing lower chl a content in the Fe‐stressed cells. Chl a per cell volume decreased by 30% as temperature decreased from 20 to 10° C under Fe‐replete conditions, but chl a decreased by 50% from Fe‐replete to Fe‐stressed conditions.     

PHYSIOLOGICAL PROFILES OF SYNECHOCOCCUS (CYANOPHYCEAE) IN IRON-LIMITING CONTINUOUS CULTURES1

We examined the physiology and biochemistry associated with the iron-limited continuous culture of the halotolerant cyanobacterium Synechococcus PCC 7002. Biomass production, photosynthetic pigment levels, photosynthetic efficiency, and the production of hydroxamate- and catechol-type siderophores are reported for cells grown over a range of available iron concentrations. The relationship between the yield of Synechococcus PCC 7002 in iron-limited chemostats and the concentration of available iron was not linear. Synechococcus PCC 7002 expressed an inducible physiological response that led to alterations either in the cellular iron quotient or, more likely, in levels of available iron due to induced iron-scavenging processes. During iron limitation these cyanobacteria produced components consistent with the activation of a high-affinity iron transport system; both hydroxamate- and catechol-type siderophores were detected. Iron-limited Synechococcus PCC 7002 also reduced CO₂ fixation rates from luxury levels to a rate that matched the cellular growth rate, presenting interesting implications for oceanic carbon flux models.     

PREDICTING THE KINETICS OF DISSOLVED INORGANIC CARBON LIMITED GROWTH FROM THE SHORT-TERM KINETICS OF PHOTOSYNTHESIS IN SYNECHOCOCCUS LEOPOLIENSIS (CYANOPHYTA)1

The blue-green alga (Cyanobacterium) Synechococcus leopoliensis (Racib.) Komarek was grown in dissolved inorganic carbon [DIC]-limited chemostats over the entire range of growth rates. At each growth rate, the kinetics of photosynthesis with respect to [DIC] and the maximal rate of photosynthesis (P_max) were determined. The half-saturation constant for [DIC]-limited photosynthesis (K_1/2^DIC) for cells growing below 1.7 d^?1 was constant (4.7 μM) whereas for growth rates between 1.7 d^?1 and 2.1 d^?1 (μ_max) the kinetics of photosynthesis were multiphasic with an apparent K_1/2^DIC between 1.5–2.0 mM. P_max increased in a linear fashion with growth rate for growth rates below 1.7 d^?1. No trend in P_max was apparent for growth rates greater than 1.7 d^?1. These kinetic parameters were used to predict a growth rate versus [DIC] relationship. Results show that the Monod relationship is a physiologically valid expression of growth as a function of [DIC] provided (K_1/2^DIC) remains constant. The major change in (K_1/2^DIC) as μ approaches μ_max results in the conclusion that two separate and distinct Monod equations must be used to describe growth as a function of DIC over the entire growth range. These results point to a major discontinuity in the μ vs. [DIC] curve at 1.7 d^?1 which corresponds to the change from high to low affinity photosynthetic kinetics. We believe these results account for the previously described deficiencies of the Monod equation in describing [DIC]-limited algal growth.     

DIFFERENCES IN GROWTH AND PHYSIOLOGY OF MARINE SYNECHOCOCCUS (CYANOBACTERIA) ON NITRATE VERSUS AMMONIUM ARE NOT DETERMINED SOLELY BY NITROGEN SOURCE REDOX STATE1

The preference of phytoplankton for ammonium over nitrate has traditionally been explained by the greater metabolic cost of reducing oxidized forms of nitrogen. This “metabolic cost hypothesis” implies that there should be a growth disadvantage on nitrate compared to ammonium or other forms of reduced nitrogen such as urea, especially when light limits growth, but in a variety of phytoplankton taxa, this predicted difference has not been observed. Our experiments with three strains of marine Synechococcus (WH7803, WH7805, and WH8112) did not reveal consistently faster growth (cell division) on ammonium or urea as compared to nitrate. Urease and glutamine synthetase (GS) activities varied with nitrogen source in a manner consistent with regulation by cellular nitrogen status via NtcA (rather than by external availability of nitrogen) in all three strains and indicated that each strain experienced some degree of nitrogen insufficiency during growth on nitrate. At light intensities that strongly limited growth, the composition (carbon, nitrogen, and pigment quotas) of WH7805 cells using nitrate was indistinguishable from that of cells using ammonium, but at saturating light intensities, cellular carbon, nitrogen, and pigment quotas were significantly lower in cells using nitrate than ammonium. These and similar results from other phytoplankton taxa suggest that a limitation in some step of nitrate uptake or assimilation, rather than the extra cost of reducing nitrate per se, may be the cause of differences in growth and physiology between cells using nitrate and ammonium.     

EFFECTS OF MODERATE HEAT STRESS AND DISSOLVED INORGANIC CARBON CONCENTRATION ON PHOTOSYNTHESIS AND RESPIRATION OF SYMBIODINIUM SP. (DINOPHYCEAE) IN CULTURE AND IN SYMBIOSIS1

The influence of temperature and inorganic carbon (C_i) concentration on photosynthesis was examined in whole corals and samples of cultured symbiotic dinoflagellates (Symbiodinium sp.) using combined measurements from a membrane inlet mass spectrometer and chl a fluorometer. In whole corals, O₂ production at 26°C was significantly limited at C_i concentrations below ambient seawater (～2.2 mM). Further additions of C_i up to ～10 mM caused no further stimulation of oxygenic photosynthesis. Following exposure to 30°C (2 d), net oxygen production decreased significantly in whole corals, as a result of reduced production of photosynthetically derived oxygen rather than increased host consumption. Whole corals maintained a rate of oxygen evolution around eight times lower than cultured Symbiodinium sp. at inorganic carbon concentrations <2 mM, but cultures displayed greater levels of photoinhibition following heat treatment (30°C, 2 d). Whole corals and cultured zooxanthellae differed considerably in their responses to C_i concentration and moderate heat stress, demonstrating that cultured Symbiodinium make an incongruous model for those in hospite. Reduced net oxygen evolution, in whole corals, under conditions of low C_i (<2 mM) has been interpreted in terms of possible sink limitation leading to increased nonphotochemical energy dissipation. The advantages of combined measurement of net gas exchange and fluorometry offered by this method are discussed.     

EFFECTS OF TEMPERATURE,SALINITY, IRRADIANCE AND DIURNAL PERIODICITY ON GROWTH AND PHOTOSYNTHESIS IN THE DIATOM NITZSCHIA AMERICANA; LIGHT-SATURATED GROWTH1

The physiological response of an estuarine clone of Nitzschia americana Fryx3ell was measured under experimental conditions of temperature and salinity which represent the average range of these variables in the Cape Fear River Estuary, North Carolina. The influence of temperature (10, 15, 20, 25, 30°C) and salinity (8, 15, 20, 26, 32‰) on specific growth rates, μ, and parameters of photosynthesis-irradiance curves, α, and P_max were measured during maximum and minimum rates of diurnal photosynthesis using axenic semi-continuous batch cultures maintained at an irradiance saturating for photosynthesis (140 μE m^-2·s-1). There was an increase in μ with increasing temperature up to a broad uptimum (25 ± 2.5°C), above which μ gradually declined. At the predicted optimum temperature of 25°C, μ increased as a linear function of salinity. oth light-limited (α) amd light-saturated (P^max) rates of photosynthesis increased as salinity decreased. The effect of temperature on a and P^max was complex and dependent on salinity. P^max exhibited a diurnal periodicity, whereas estimates of a were not significantly different between sampling periods. Growth efficiencey opf N. americana, calculated as the ratio between specific growth rates and rates of gross photosynthesis, increased with an increase in salinity with a maximum at the predicted optimum temperature and salinity of 25°C and 32‰, suggesting and uncoupling between photosynthesis and growth at nonoptimum growth conditions.     

CYANOBACTERIAL ASSEMBLAGES IN PERMANENT ICE COVERS ON ANTARCTIC LAKES: DISTRIBUTION, GROWTH RATE, AND TEMPERATURE RESPONSE OF PHOTOSYNTHESIS

The proliferation of microalgae in the McMurdo Dry Valleys of Antarctica is intricately linked to the seasonal cycle involving the freezing and melting of water. Anecdotal observations and preliminary sampling have found cyanobacterial cells in ice covers on lakes in the McMurdo Dry Valleys, and several of these ice covers are known to undergo seasonal freeze–thaw cycles. Therefore, we sought to determine the distribution and abundance of cyanobacterial assemblages in several permanent ice covers throughout the McMurdo Dry Valleys and to determine their rates of growth and their photosynthetic physiologies upon encountering liquid water. We found that the majority of the permanent ice covers contained cyanobacterial assemblages in close association with sedimentary material. Cyanobacterial biomass was conspicuously absent in sediment-free ice covers, suggesting that the seasonal interaction between the sediments, ice, and solar radiation present the necessary liquid water environment for cyanobacterial growth. All assemblages exhibited extremely low rates of photosynthesis when first exposed to liquid water. Despite the low rates of photosynthesis, a large proportion (41%) of the photosynthate was incorporated into protein, indicating that the cells were undergoing efficient net cellular growth. The short-term response (24 h) of photosynthesis to a range of temperatures showed optimum rates occurring at temperatures >15° C, which is similar to those of psychrotrophic cyanobacteria isolates from soil and stream habitats, which we believe provides the inoculum for the in- ice habitats.     

EFFECTS OF TEMPERATURE ON GROWTH,LIGHT ABSORPTION,AND QUANTUM YIELD IN DUNALIELLA TERTIOLECTA (CHLOROPHYCEAE)1

The effects of growth temperature on the marine chlorophyte Dunaliella tertiolecta Butcher were studied to provide a more mechanistic understanding of the role of environmental factors in regulating bio-optical properties of phytoplankton. Specific attention was focused on quantities that are relevant for modeling of growth and photosynthesis. Characteristics including chlorophyll a (chl z)-specific light absorption (a*_ph(λ)), C:chl a ratio, and quantum yield for growth (φ_μ) varied as functions of temperature under conditions of excess light and nutrients. As temperature increased over the range examined (12°-28°C), intracellular concentrations of chl a increased by a factor of 2 and a*_ph(λ) values decreased by more than 50% at blue to green wavelengths. The lower values of a*_ph(λ) were due to both a decrease in the abundance of accessory pigments relative to chl a and an increase in pigment package effects arising from higher intracellular pigment concentrations. Intracellular pigment concentration increased as a consequence of higher cellular pigment quotas combined with lower cell volume. At high growth temperatures, slightly more light was absorbed on a per-cell-C basis, but the dramatic increases in growth rate from μ= 0.5 d^?1 at 12° C to μ= 2.2 d^?1 at 28°C were primarily due to an increase in φ_μ (0.015–0.041 mol C (mol quanta)^?1). By comparison with previous work on this species, we conclude the effects of temperature on a*_ph(λ) and φ_μ are comparable to those observed for light and nutrient limitation. Patterns of variability in a*_ph(λ)and φ_μ as a function of growth rate at different temperatures are similar to those previously documented for this species grown at the same irradiance but under a range of nitrogen-limited conditions. These results are discussed in the context of implications for bio-optical modeling of aquatic primary production by phytoplankton.     

COMBINED EFFECTS OF ULTRAVIOLET RADIATION AND TEMPERATURE ON MORPHOLOGY,PHOTOSYNTHESIS, AND DNA OF ARTHROSPIRA (SPIRULINA) PLATENSIS (CYANOPHYTA)1

Natural levels of solar UVR were shown to break and alter the spiral structure of Arthrospira (Spirulina) platensis (Nordst.) Gomont during winter. However, this phenomenon was not observed during summer at temperatures of ～30°C. Since little has been documented on the interactive effects of solar UV radiation (UVR; 280–400 nm) and temperature on cyanobacteria, the morphology, photosynthesis, and DNA damage of A. platensis were examined using two radiation treatments (PAR [400–700 nm] and PAB [PAR + UV‐A + UV‐B: 280–700]), three temperatures (15, 22, and 30°C), and three biomass concentrations (100, 160, and 240 mg dwt [dry weight] · L^?1). UVR caused a breakage of the spiral structure at 15°C and 22°C, but not at 30°C. High PAR levels also induced a significant breakage at 15°C and 22°C, but only at low biomass densities, and to lesser extent when compared with the PAB treatment. A. platensis was able to alter its spiral structure by increasing helix tightness at the highest temperature tested. The photochemical efficiency was depressed to undetectable levels at 15°C but was relatively high at 30°C even under the treatment with UVR in 8 h. At 30°C, UVR led to 93%–97% less DNA damage when compared with 15°C after 8 h of exposure. UV‐absorbing compounds were determined as negligible at all light and temperature combinations. The possible mechanisms for the temperature‐dependent effects of UVR on this organism are discussed in this paper.     

RELATIONSHIP BETWEEN PHOTOSYNTHETIC OXYGEN CYCLING AND CARBON ASSIMILATION IN SYNECHOCOCCUS WH7803 (CYANOPHYTA)1

Mass spectrometric analysis of oxygen uptake and evolution in the light by marine Synechococcus WH7803 indicated that the respiration rate was near zero at low irradiance levels but increased significantly at high irradiances. The light intensity (I_r) at which oxygen uptake began to increase with increasing light intensity depended on the growth irradiance of the culture. In each case, I_r coincided with the minimum light intensity for saturation of carbon assimilation (I_k). At irradiances >I_r, net oxygen evolution rates paralleled carbon assimilation rates. Oxygen uptake at high light intensities was inhibited by DCMU, indicating that oxygen uptake was due to Mehler reaction activity. The onset of Mehler activity at I_k supports the idea that oxygen becomes an alternative sink for electrons from photosystem I when NADPH turnover is limited by the capacity of the dark reactions to utilize reductant.     

CO2浓度变化对拟柱胞藻生长与光合作用的影响

为了探讨大气CO₂浓度升高对水华藻类的影响,利用水华蓝藻-拟柱胞藻作为实验材料,研究了CO₂浓度升高对其生长生理和光合作用的影响,结果表明CO₂浓度升高,导致拟柱胞藻的生物量、最大光合放氧速率、光合效率显著增加。当CO₂浓度为700 mg/L以下,暗呼吸速率和光饱和点无明显影响,而CO₂浓度为1000 mg/L时,暗呼吸速率和光饱和点显著提高。随着CO₂浓度增加,藻细胞光合作用对无机碳的亲和力降低,同时胞外碳酸酐酶活性显著下降。这表明大气CO₂浓度的增加,有利于拟柱胞藻的生长和光合,进而增加了水华发生的风险。     

ADAPTATION OF PHOTOSYNTHESIS IN LAMINARIA SACCHARINA (PHAEOPHYTA) TO CHANGES IN GROWTH TEMPERATURE1

The effect of growth temperature on photosynthetic metabolism was studied in the kelp Laminaria saccharina (L.) Lamour. Photosynthesis was subject to phenotypic adaptation, with almost constant photosynthetic rates being achieved at growth temperatures between 0 and 20° C. This response involved: (1) an inverse relationship between growth temperature and photosynthetic capacity, (2) a reduction in the Q₁₀ value for photosynthesis of L. saccharina grown at 0 and 5° C compared with 10, 15 and 20° C grown sporophytes, and (3) an acquired tolerance of photosynthesis to temperatures between 15–25° C (which inhibited photosynthesis in 0 and 5° C grown L. saccharina) in sporophytes grown at 10, 15 and 20° C. The physiological basis of these adaptations is discussed in terms of observed changes in activities and kinetics of the Calvin cycle enzyme ribulose-1, 5-bisphosphate carboxylase (oxygenase) and efficiency of light harvesting-electron transport systems.     

EFFECTS OF LIGHT INTENSITY AND TEMPERATURE INTERACTIONS ON GROWTH CHARACTERISTICS OF PHAEODACTYLUM TRICORNUTUM (BACILLARIOPHYCEAE)1

Cell division rate, carbon fixation per cell, cell width and chloroplast length of Phaeodactylum tricornutum Bohlin were determined at 30 different combinations of light intensity and temperature. Division rate peaked at 23° C or less depending on light intensity. For each light intensity studied, carbon fixation increased directly with growth temperature from 14 to 25° C. The slope of this relationship was modified by light intensity. Cells grown at 23–25° C tended to be larger than those grown at lower temperatures, possibly due to increased carbon fixation per cell coupled with lower division rates. Chloroplasts were largest at a combination of temperatures above 21° C and low light intensities. This effect could cause cells to sink at a higher than normal rate due to reduced vacuole size and is presented as a possible mechanism affecting the distribution of P. tricornutum.     

EFFECTS OF ULTRAVIOLET RADIATION ON PHOTOSYNTHESIS IN THE SUBTROPICAL MARINE DIATOM,CHAETOCEROS GRACILIS (BACILLARIOPHYCEAE)1

Acclimation to ambient ultraviolet radiation (UVR) was examined in a subtropical marine diatom, Chaetoceros gracilis Schutt. Short-term exposure to UVR (<24 h) reduced the efficiency of photosynthetic energy conversion, carbon fixation, activity of 1,5-bisphosphate carboxylase-loxygenase (RUBISCO), and the rapid turnover of the putative Dl reaction center (32 kda) protein, whereas longer-term exposure to ambient UVR (24–48 h) revealed a steady-state acclimation, defined as recovery of carbon fixation and RUBISCO activity to rates equivalent to treatments without exposure to UVR. The turnover of D1 and chlorophyll a (Chl a) remained high during exposure to UVR. Efficiency of energy conversion by photosystem II, measured with double flash (pump and probe) fluorometry, increased by 24% in cells acclimated to UVR. Acclimation to UVR had no detectable effect on the functional absorption cross-section or cellular concentrations of Chl a, Chl c, or total carotenoids. However, the maximum rate of carbon fixation was reduced by UVR on a Chl a basis but remained unaffected on a per-cell basis. Response to UVR exposure in this subtropical diatom has two components: a short-term inhibitory response and a longer-term acclimation process that ameliorates the inhibition of carbon fixation.     

THE EFFECT OF IRON NUTRITION ON PHOTOSYNTHESIS AND NITROGEN FIXATION IN CULTURES OF TRICHODESMIUM (CYANOPHYCEAE)1

Cultures of Trichodesmium NIBB 1067 were grown in the synthetic medium AQUIL with a range of iron added from none to 5 × 10^?7 M Fe for 15 days. Chlorophyll-a, cell counts, and total cell volume were two or three times higher in medium with 10^?7 M Fe than with no added Fe. Oxygen production rate per chlorophyll-a was over 60% higher with higher iron. Increased iron stimulated photosynthesis at all irradiances from about 12–250 μE · m^?2· s^?1. Nitrogen fixation rate, estimated from acetylene reduction, for 10^?7 and 10^?8 M Fe cultures was approximately twice that of the cultures with no added Fe. The range of rates of O₂ production and N₂ fixation in cultures at the iron concentrations we used were similar to the rates from natural samples of Trichodesmium from both the Atlantic, and the Pacific oceans. This similarity may allow this clone to be used, with some caution, for future physiological ecology studies. This study demonstrates the importance of iron to photosynthesis and nitrogen fixation and suggests that Trichodesmium plays a central role in the biogeochemical cycles of iron, carbon and nitrogen.     

PHOTOINHIBITION OF PHOTOSYNTHESIS IN ASTERIONELLA FORMOSA (BACILLARIOPHYCEAE)1, 2

A laboratory study was made of the inhibition of photosynthesis, as measured by the radiocarbon method, of Asterionella Formosa Hass. by light from tungsten halogen lamps of irradiances up to that of full sunlight. The observed inhibitions were of total fixation of carbon and were not due to greatly increased release of extracellular products of photosynthesis. Inhibition increased width irradiance and with time of exposure. It was greater at high temperature, at high oxygen concentrations and when the cells were nutrient deficient. Recovery from exposure to high irradiances took place both in the dark and at low irradiances. The inhibition, which is similar to that observed under natural conditions, has the characteristics of photooxidative damage to both photochemical and dark reaction mechanisms.