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     

Primary production as influenced by diel periodicity of phytoplankton photosynthesis

Diel periodicity in parameters of photosynthesis-irradiance (P-I) curves was incorporated into calculations of integral daily phytoplankton production for the Santa Barbara Channel off southern California (USA). Model equations of the relationship between photosynthesis and light were used in combination with observed slope () and asymptote (P _max) values presented in the preceding paper. Primary production was always 19 to 39% less than comparable estimates obtained with the assumption of constant maximum daily and P _max values. Regardless of which P-I formulation was used of 6 tested, observed production (using a temporal series of simulated in situ incubations) ranged from 13% less to 25% more than estimates from constant midday and P _max values. The amplitude and timing of diel oscillations differed somewhat among 3 field stations. Maximum to minimum ratios ranged from approximately 3 to 5 for , and 4 to 6 for P _max. The differences in amplitude and timing of oscillations in P-I curves both contributed to errors in calculating phytoplankton production. Thus, photosynthetic periodicity in the upwelling area of the Santa Barbara Channel influences phytoplankton production. There were oscillations in both and P _max, and the time-dependence of these parameters should be considered to improve the accuracy of predictive models of primary productivity.     

Short-term variability of photosynthesis in natural marine phytoplankton populations

Harmonic regression analysis has been used to determine the short-term variability in the photosynthetic rate (mgC/mg chlorophyll a/h) of phytoplankton in three inlets of Japan. In natural water without large zooplankton present, the photosynthetic rate [log P=log (100xmgC/mg chlorophyll a/h)] can be expressed as (B+A cos T). Factor B represents the average photosynthetic rate, of which the maximum is usually designated as P _max, and Factor A corresponds to the slope of the regression line. The phase of the periodicity, represented by T, is adjusted to give the highest correlation: usually T is expressed as [360/24 x (local time + 4)] in degrees. The correlation between Factors A and B is very high (r=0.95, P<0.001), indicating that Factor A may depend upon Factor B (potential activity of chlorophyll a). Both Factors A and B decrease with decreasing irradiance, but the slope of each regression between Factor A and irradiance varies with season. Continuous darkness reduces the phase of the periodicity to one cycle a day when phytoplankton has multiple cycles of photosynthetic rate per day. Adequate nutrient supply from zooplankton regeneration may cause an increase in Factor B; however, excess density of zooplankton decreases Factor A.     

The effects of three oils on marine phytoplankton photosynthesis

The effects of 3 oils (Venezuelan crude, No. 2 fuel, and No. 6 fuel) on the photosynthesis of natural phytoplankton communities from Bedford Basin, Nova Scotia (Canada), and the northwest Atlantic Ocean between Halifax and Bermuda were examined using a radiocarbon method. The 3 oils can inhibit photosynthesis, and the degree of inhibition depends upon oil type and concentration. The No. 2 fuel oil was the most toxic. Under certain conditions, low concentrations of Venezuelan crude oil can stimulate photosynthesis. On the basis of these results, it is concluded that present levels of oil contamination in Bedford Basin could be inhibiting photosynthesis by a few percent, while present levels in open ocean water have no apparent deleterious effect on photosynthesis.Bedford Institute of Oceanography Contribution.     

Products of photosynthesis in natural populations of marine phytoplankton from the Gulf of Maine

We have measured the photosynthetic assimilation of ¹⁴C-carbon dioxide into (1) ethanol-soluble, (2) hot-trichloroacetic acid (TCA)-soluble (polysaccharide), and (3) protein fractions of natural populations of marine phytoplankton. Diurnal studies showed a continuing incorporation of carbon-14 into the protein fraction during hours of darkness. This was accompanied by a concomitant decrease in the proportion assimilated into polysaccharide. When incorporation was measured under constant experimental conditions, the pattern of photosynthesis did not vary from one time of day to another. At one station approximately 12 km south of Boothbay Harbor, the proportion of carbon entering protein showed marked seasonal changes. During the winter, approximately 10 to 20% of the fixed carbon was incorporated into protein. During the summer the value increased to 22 to 35%. Between these times, a transient high value of 37 to 47% of the fixed carbon entering protein coincided with the spring bloom. The increases in proportion incorporated into protein were largely paralleled by equivalent decreases in the polysaccharide fraction. The proportion of carbon incorporated into protein during photosynthesis also increased markedly at reduced light intensities. This increase occurred both when populations were incubated in neutral-density filters and when incubated at increasing depths in the photic zone. There was little consistent and significant difference between the neutral-density filters and depth in the water column, suggesting a minimal role for light quality. The extent of the increased relative rate of protein synthesis at the lower light intensities depended on the nutritional state of the phytoplankton. For example, summer populations from water containing low concentrations of inorganic nutrients responded less dramatically to reduced light intensities than did populations from nutrient-rich waters.     

Effects of light intensity on photosynthesis and dark respiration in six species of marine phytoplankton

Using an oxygen polarographic electrode, the shapes of photosynthetic curves and the effects of light on dark respiration in 6 species of marine phytoplankton wer examined. The species used were Skeletonema costatum, Ditylum brightwellii, Cyclotella nana (Thalassiosira pseudonana) (all Bacillariophyceae), Dunaliella tertiolecta (Chlorophyceae), Isochrysis galbana (Haptophyceae), and Gonyaulax tamarensis (Dinophyceae). A hysteresis was observed in all species examined with respect to increasing and decreasing light. Compensation light intensities varied by over 4 orders of magnitude, suggesting that the 1% light depth is an ambiguous measure of the euphotic zone. The data suggest that dark respiration accounts for ca. 25% of gross photosynthesis, but is species-dependent. In addition, respiration versus cell size does not describe an inverse exponential function over the size scales examined.This research was performed under the auspices of the United States Energy Research and Development Administration under Contract No. EY-76-C-02-0016.     

Polychlorinated biphenyl (PCB) effects on marine phytoplankton photosynthesis and cell division

Inhibition of photosynthesis and cell division by polychlorinated biphenyls (PCBs) was studied using 7 marine phytoplankton species representing 4 algal classes. PCB concentrations as low as 1.0 g l^-1 reduced cell division of Thalassiosira pseudonana 3H and Isochrysis galbana. Both photosynthesis and cell division of T. pseudonana 3H, Chaetoceros socialis, Skeletonema costatum, T. pseudonana 13-1, Monochrysis, lutheri and I. galbana were inhibited at a PCB concentration of 10.0 g l^-1. The effects on photosynthesis were immediate and probably resulted in reduced rates of cell division. Interspecific differences in susceptibility were observed. These differences have significance with respect to primary production and the species composition of phytoplankton communities. The initial slopes of photosynthesis-irradiance (P-I) curves for the diatoms S. costatum and T. pseudonana 3H were reduced in the presence of PCBs. These results suggest that PCBs affect the photosynthetic light reactions.     

Diel changes in phytoplankton photosynthetic efficiency in Brackish waters

From 18 to 23 September 1974, investigations on the diel changes in phytoplankton were carried out in the Baltic Sea. Every 4 h, water samples were collected from 2 and 15 m, and PO₄, chlorophyll a, temperature, salinity, pH, phytoplankton composition and phytoplankton light photosynthesis relationship were determined. Continuous measurements of surface irradiance and some estimations of zooplankton were also made. P ^B (photosynthesis per unit chlorophyll a at low light levels of 2·10^-2 cal cm^-2 min^-1) revealed only random variation during the sampling period, i.e., 1.0 to 1.6 mg C (mg chlorophyll a)^-1 h^-1. P _m ^B (Light-saturated photosynthesis per unit of chlorophyll a) displayed pronounced diel fluctuations with the highest value of about 6 mg C (mg chlorophyll a)^-1 h^-1 around noon, and the lowest value of about 2.5 mg C (mg chlorophyll a)^-1 h^-1 during the night, during which latter period the value of P _m ^B was more or less constant. Reasons for the diel fluctuations are discussed, and an equation which describes these fluctuations is proposed. Using this equation, the daily phytoplankton production estimated in incubators by a previously described method can be corrected for the time of day at which samples are collected.     

Hydrocarbons of marine phytoplankton

The hydrocarbon contents of 23 species of algae (22 marine planktonic), belonging to 9 algal classes, were analyzed. The highly unsaturated 3,6,9,12,15,18-heneicosalhexaene predominates in the Bacillariophyceae, Dinophyceae, Cryptophyceae, Haptophyceae and Euglenophyceae. Rhizosolenia setigera contains n-heneicosane, presumably derived from the hexaolefin by hydrogenation. Two isomeric heptadecenes have been isolated: the double bond is located in 5-position in the bluegreen alga Synechococcus bacillaris and in 7-position in 2 green algae. Our complete analyses are discussed in the context of earlier data; some generalizations appear no longer valid. Hydrocarbon analysis of marine algae should provide a tool for the investigation of the dynamics of the marine food chain. Knowledge now available provides the background needed for distinguishing between hydrocarbons of recent biogenic origin and hydrocarbon pollutants from fossil fuels.Contribution No. 2576 of the Woods Hole Oceanographic Institution, USA.     

The effect of intensity and quality of illumination on the photosynthesis of some tropical marine phytoplankton

Using solar energy as a source of illumination, photosynthesis in 11 species of marine plankton algae was studied as a function of light intensity. From the photosynthesis-light curve for each organism, the saturation points (I _k ) in different organisms were determined. Among the diatoms and flagellates, the highest I _k (saturation point) values were found in Rhizosolenia styliformis and Dinophysis miles respectively. When the organisms were exposed to a portion of the visible spectrum starting from 700 m, the photosynthesis was found to be related to the radiant energy. The missing portions of the spectrum produced no significant change in the rate of photosynthesis. The photosynthetic response shown by the different organisms was strikingly similar, which signifies that, despite the qualitative dissimilarities which the organisms may possess in their pigment composition, they are capable of much chromatic adaptation.     

Products of photosynthesis by marine phytoplankton: the effect of environmental factors on the relative rates of protein synthesis

A method is presented by which the gross pattern of photosynthetic carbon-dioxide fixation in marine phytoplankton can be determined. It depends on differential solvent extraction yielding an ethanol-soluble, a hot TCA-soluble (polysaccharide) and a residue (protein) fraction. Using this fractionation technique, the effects of various environmental factors on the pattern of photosynthesis by the marine diatom Phaeodactylum tricornutum (Bohlin) have been investigated. Low light intensities and increasing degrees of nitrogen limitation in a chemostat increase markedly the relative rates of protein synthesis. Growth of the alga at lower temperatures also increases the proportion of carbon incorporated into the protein fraction. This increased protein syntheses is generally at the expense of the polysaccharide fraction. Preliminary experiments have established the suitability of this fractionation method for natural populations of phytoplankton and have shown similar effects of light intensity on the relative rates of protein synthesis.     

Diel oscillations of the photosynthesis-irradiance (P-I) relationship in natural assemblages of phytoplankton

Diel oscillations in the photosynthesis-irradiance (P-I) relationship are described for marine phytoplankton assemblages at 6 stations in an upwelling area off the southern California coast (USA) between May and August 1980. The initial slope () and asymptote (P _max) of P-I curves changed significantly over the day; both parameters were in phase and had similar changes in amplitude. The diel oscillations in photosynthesis appeared unrelated to changes in chlorophyll a concentrations. Amplitudes of daily variations in photosynthesis ranged from approximately 3 to 9, as measured by the maximum to minimum ratio for photosynthetic capacity (P _max). Diatom-rich samples collected during an upwelling event and those dominated by dinoflagellates both had midday to early afternoon maxima in and P _max. Samples from other locations had peak photosynthetic activity later in the afternoon. The relationship between and P _max was consistent in all phytoplankton samples analyzed, with a surprisingly high correlation considering the spatial and temporal scales encompassed in this study. These results indicate that the photosynthesis-irradiance (P-I) relationship is time-dependent and, moreover, that changes in and P _max are closely coupled for a variety of natural phytoplankton assemblages.     

Cycling of selenite and selenate in marine phytoplankton

The marine phytoplankton Dunaliella tertiolecta, Cachonina niei, Thalassiosira nordenskioldii, Phaeodactylum tricornutum, and Chaetoceros sp. were incubated with a range of molar concentrations of sodium-selenite (Na₂-SeIVO₃) and sodium-selenate (Na₂-SeVIO₄) to examine further their role in metabolic cycling of selenium in ocean waters. At low selenium concentrations, approaching those found naturally in seawater (10^-10 to 10^-9 M), all species distinguished between selenite and selenate, and actively concentrated selenite from the incubating medium while only marginally accumulating selenate. At much higher concentrations (10^-8 to 10^-6 M), selenate was also taken up. At the highest concentration tested, i.e., 10^-5 M with C. niei, after an immediate rapid uptake in the first 24 h, the intracellular selenite and selenate levels dropped to about 35 to 50% of the initial peak values. These observations suggest an uptake mechanism in these algae which, at normal ambient concentrations of selenium (10^-9 M), preferentially selects selenite and excludes selenate. At much higher concentrations (10^-8 M), the mechanism becomes overloaded and both selenium species enter the cells. Intracellularly, selenite became associated primarily with protein and amino acid fractions, in approximately equal proportions, while only ca. 4% of total intracellular selenium was found in the lipid fraction. Trace amounts of selenate that entered the cells, mainly during the first minutes of exposure, also entered the protein and amino acid components, but over time were increasingly associated with the protein fraction only. At the end of a 10-d incubation of algal cells in selenite-spiked medium, less than 25% of total Se in the medium could in fact be identified analytically as selenite. This suggests the presence of a non-selenite metabolite, possibly released back into the medium from the algae.     

Light stimulation of phosphate uptake in marine phytoplankton

A phosphate uptake system responding positively to light would offer the algae a competitive advantage over bacteria, particularly when phosphorus is limiting. Effects of light on phosphate uptake in phytoplankton were investigated in view of conflicting literature reports, to test the hypothesis that both the P status and the light history of the population controlled the response. Maximum stimulation, to about 2.7x dark uptake, was observed at a Strait of Georgia, British Columbia, Canada station where phytoplankton was phosphorus sufficient but light limited. Stimulation was similar, about 1.8 x the dark rate, in the Northern Sargasso Sea, where phytoplankton was probably limited by phosphate but not by light. Minimal stimulation was observed in the Sheepscot Estuary, Maine, USA, where the population appeared to be neither phosphorus nor light limited.     

Measurements of electron transport activities in marine phytoplankton

A simple and sensitive spectrophotometric method of measuring electron transfer in the transport system (ETS) in marine phytoplankton has been developed and characterized. The assay is based on the reduction of the tetrazolium salt 2-(p-iodophenyl)-3-(p-nitrophenyl)-5-phenyl tetrazolium chloride (INT) by homogenates in the presence of the nonionic detergent Triton X-100. The simplicity and sensitivity of this assay have considerable advantage over existing methods of measuring ETS activity in marine samples where numerous assays must be rapidly carried out on samples which often demonstrate low activities. Several established substrates and inhibitors of electron transport in the ETS were investigated, and each is discussed in relation to the site of INT reduction in the diatoms Skeletonema costatum and Chaetoceros debilis and in the unicellular green alga Dunaliella tertiolecta. The results from these studies suggest that the maximum rate of electron transport, V _max, is measured in each case.     

Decomposition of urea associated with photosynthesis of phytoplankton in coastal waters

Decomposition of urea in seawater was studied in Mikawa Bay, a shallow eutrophic bay on the southern coast of central Japan. The urea concentration in seawater ranged from 1.3 to 5.9 μg-at. N/1 and comprised 12 to 40% of the dissolved organic nitrogen. Using ¹⁴C labelled urea, the rate of CO₂ liberation from urea and the incorporation rate of urea carbon into the particulate organic matter were determined. For the surface samples, high rates of CO₂ liberation from urea as well as the incorporation of urea carbon into the particulate organic matter were observed in the light, while much lower rates were obtained in the dark. Incubation experiments with exposure to different light intensities revealed that the rate of CO₂ liberation from urea and the incorporation of urea carbon into particulate organic matter changed with light intensity, showing a pattern similar to that of photosynthesis. The highest liberation and incorporation rates were observed at 12,000 lux. Incubation in light and in dark produced marked decreases and increases, respectively, in urea and ammonia, while no appreciable changes were observed for nitrate and nitrite. It is suggested that urea decomposition associated with photosynthetic activity of phytoplankton is one of the major processes of urea decomposition, and that it plays a significant role in the nitrogen supply for phytoplankton in coastal waters.     

Buoyancy of natural populations of marine phytoplankton

Buoyancy of natural populations of marine phytoplankton was studied in a fjord in western Norway during the diatom bloom and in autumn. The study was carried out under approximate in situ conditions by means of an apparatus described in the paper. During the spring bloom, positive buoyancy was observed only once. Sinking rates of individual fractions ranged from 0 to more than 9 m day^-1, and the mean sinking rates of the total chlorophyll content from 0 to at least 2. 2 m day^-1. The highest rates occurred in the post-bloom period, while sinking appeared negligible from the onset of the bloom up to its culmination. In autumn, the population was dominated by small, flagellated cells. Positive buoyancy, or upward migration, was then observed in two out of three experiments.     

Effects of diurnal variations in phytoplankton photosynthesis obtained from natural fluorescence

Effects of diurnal variation in phytoplankton photosynthesis on estimating daily primary production (DPP) were examined using field data from Sagami Bay, Japan. DPP at 5 m depth was calculated from the continuous data of chlorophyll a (Chl a) and light intensity monitored by a natural fluorescence sensor with and without considering time-dependent changes in the photosynthesis–irradiance (P–E) relationship. Chl a could be estimated from natural fluorescence examining the variations in the quantum yield of fluorescence (φ _f) and Chl a-specific light absorption coefficient (a*_ph), and relating them to Chl a. The P–E relationship was determined by water sampling three times daily. A distinct diurnal pattern was observed for the maximum photosynthetic rate (P*_max), being maximal at noon, while periodicity of the maximum light utilization coefficient (α*) was less obvious. The actual DPP was calculated by interpolating the P–E parameters from those obtained at dawn, noon, and dusk. For comparison, DPP was calculated by fixing the P–E parameters as the constants measured at dawn, noon or dusk for a day. The difference from the actual DPP was small when the P–E parameters measured at dawn (3% on average) and noon (5%) were used as the constants for a day. The difference was largest when the values at dusk were used (−43%). The medium values of P*_max at dawn, its low values at dusk, and the fact that a major part of the DPP was produced around noon were responsible for these results. The present study demonstrates that measurement of the P–E parameters at dawn or noon can give a good estimation of DPP from natural fluorescence.     

Size structure of phytoplankton biomass and photosynthesis in subtropical Hawaiian waters

In a subtropical Hawaiian ecosystem, phytoplanton size structure analyses (November–December, 1980) showed that ultraplankton (>3μm), nanoplankton (>20μm) and netplankton (>20μm) accounted for ca. 80, 98, and 2% of total chlorophyll standing stock, respectively, on the basis of chlorophyll. Similar trends were evident, for other biomass indices (e.g. cell numbers, total cell volume, ATP, particulate organic carbon, particulate organic nitrogen). The ultraplankton fraction consisted primarily of small flagellates (1 to 3 μm diam) and coccoid cells (?1 μm diam); the 3 to 20 μm fraction was represented by dinoflagellates, coccolithophores, diatoms, and chrysophytes; and the netplankton fraction consisted principally of dinoflagellates and centric diatoms. Community photosynthesis had a size distribution similar to that of biomass. Sinking rates for the 3 μm, 3 to 20 μm, and >20 μm fractions averaged 0.0, 0.09, and 0.29m d^?1, respectively. The absence of measurable sinking rates for the ultraplankton, together with the relative abundance of biomass in this fraction, result in very small phytoplankton losses due to sinking in such subtropical surface waters.     

Iron-mediated effects on nitrate reductase in marine phytoplankton

The potential activity of nitrate reductase was determined in uni-algal cultures in the laboratory and in natural marine phytoplankton assemblages. In the laboratory bioassays, distinct differences in nitrate reductase activity were observed in iron replete versus depleted cultures for Emiliania huxleyi, Isochrysis galbana and Tetraselmis sp. Cells from iron-depleted cultures had 15 to 50 percent lower enzyme activity than those from iron-replete cultures. Upon addition of iron, nitrate reductase activity was enhanced in depleted cells up to levels comparable to those of the replete cells. Bioassays in the northern North Sea conducted in 1993, under low iron conditions, demonstrated similar results. Upon addition of 2.5 nM iron, a distinct enhancement, to a maximum of three times, of nitrate reductase activity was observed within 32 h after addition. Therefore, iron can stimulate nitrate reductase activity. In spite of the clean techniques used, some nitrate reductase activity was always observed. Iron deficiency was shown to impair nitrate reductase activity, but it is unlikely that nitrate reduction would cease completely.