Contribution ofSynechococcus spp. to size-fractioned primary productivity in three water masses in the Northwest Atlantic Ocean

The distribution of phycoerythrin-richSynechococcus spp. relative to eukaryotic algae and the contribution ofSynechococcus spp. toin situ primary production were compared at a neritic front, in warm-core eddy 84-E, and at Wilkinson's Basin, during a cruise to the Northwest Atlantic Ocean in July/August 1984. Immunofluorescence analyses ofSynechococcus strains demonstrated the restricted distribution of the tropical oceanic serogroup to the warm-core eddy, while strains of the neritic serogroup and those labelled by antiserum directed against a motile strain, were abundant in all three water masses. Although the majority ofSynechococcus spp. cells were observed in the 0.6 to 1 m fraction, an increasing proportion of the totalSynechococcus spp. cells were found in the 1 to 5 m fraction as nitrate concentrations increased near the base of the thermocline. From immunofluorescence analyses, we determined that the increasing proportion of largerSynechococcus spp. cells at depth was not the result of a change in strain composition, and may therefore be associated with increasing cell volume due to the enhanced nutrient supply. The contribution of the different size fractions to the total standing crop of chlorophyll and thein situ rate of photosynthesis was distincty different for the three water masses. At the neritic front, the larger photoautotrophs of the 1 to 5 m and >5 m fractions were the major contributors to chlorophyll concentrations and primary production.Synechococcus spp. appeared to provide only 6% of the dawn-to-duskin situ primary production at the neritic front. In modified Sargasso water in the warm-core eddy,Synechococcus spp. contributed 25% to thein situ rate of integrated primary production. In this warm-core eddy, the 0.2 to 0.6 m fraction made a major contribution to the standing crop of chlorophyll and primary production that equalled or exceeded that of the larger sze categories. Furthermore, at the bottom of the euphotic layer, eukaryotes numerically dominated the 0.2 to 0.6 m fraction, which contributed 61% of the primary productivity. At Wilkinson's Basin, theSynechococcus spp.-dominated 0.6 to 1.0 m fraction made the greatest contribution to the standing crop of chlorophyll an primary production, while smaller photoautotrophs (0.2 to 0.6 m) accounted for little of the chlorophyll or photosynthetic rates measured over the euphotic layer. Largest numbers ofSynechococcus spp. (2.9x10⁸ cells l^-1) occurred at the 18% isolume, coincident with a shoulder in the chlorophyll fluorescence profile and the site of maximumin situ primary productivity. At Wilkinson's Basin,Synechococcus spp. contributed 46% to thein situ photosynthesis integrated over the water-column.     

Diurnal patterns in photosynthetic capacity and depth-dependent photosynthesis-irradiance relationships inSynechococcus spp. and larger phytoplankton in three water masses in the Northwest Atlantic Ocean

The photosynthetic characteristics of prokaryotic phycoerythrin-rich populations of cyanobacteriaSynechococcus spp. and larger eukaryotic algae were compared at a neritic frontal station (Pl), in a warm-core eddy (P2), and at Wilkinson's Basin (P3) during a cruise in the Northwest Atlantic Ocean in the summer of 1984.Synechococcus spp. numerically dominated the 0.6 to 1 m fraction, and to a lesser extent the 1 to 5 m size fractions, at most depths at all stations. At P2 and P3, all three size categories of phytoplankton (0.6 to 1 m, 1 to 5 m, and >5 m) exhibited similar depth-dependent chages in both the timing and amplitude of diurnal periodicities of chlorophyllbased and cell-based photosynthetic capacity. Midday maxima in photosynthesis were observed in the upper watercolumn which damped-out in all size fractions sampled just below the thermocline. For all size fractions sampled near the bottom of the euphotic zone, the highest photosynthetic capacity was observed at dawn. At all depths, theSynechococcus spp.-dominated size fractions had lower assimilation rates than larger phytoplankton size fractions. This observation takes exception with the view that there is an inverse size-dependency in algal photosynthesis. Results also indicated that the size-specific contribution to potential primary production in surface waters did not vary appreciably over the day. However, estimates of the percent contribution ofSynechococcus spp. to total primary productivity in surface waters at the neritic front were significantly higher when derived from short-term incubator measurements of photosynthetic capacity rather than from dawn-to-duskin situ measurements of carbon fixation. The discrepancy was not due to photoinhibitory effects on photosynthesis, but appeared to reflect increased selective grazing pressure onSynechococcus spp. in dawn-to-dusk samples. Low-light photoadaptation was evident in analyses of the depth-dependency ofP-I parameters (photosynthetic capacity,P _max; light-limited slope, alpha;P _max alpha,I _k ; light-intensity beyond which photoinhibition occurs,I _b ) of the > 0.6 m communities at all three stations and was attributable to stratification of the water column. There was a decrease in assimilation rates andI _k with depth that was associated with increases in light-limited rates of photosynthesis. No midday photoinhibition ofP _max orI _b was observed in any surface station. Marked photoinhibition was detected only in the chlorophyll maximum at the neritic front and below the surface mixed-layer at Wilkinson's Basin, where susceptibility to photoinhibition increased with the depth of the collected sample. The 0.6 to 1 m fraction always had lower light requirements for light-saturated photosynthesis than the > 5 m size fraction within the same sample. Saturation intensities for the 1 to 5 m and 0.6 to 1 m size fractions were more similar whenSynechococcus spp. abundances were high in the 1 to 5 m fraction. The > 5 m fraction appeared to be the prime contributor to photoinhibitory features displayed in mixed samples (> 0.6 m) taken from the chlorophyll maxima. InSynechococcus spp.-dominated 0.6 to 1 and 1 to 5 m size fractions, cellular chlorophylla content increased 50- to 100-fold with depth and could be related to increases in maximum daytime rates of cellularP _max at the base of the euphotic zone. Furthermore, the 0.6 to 1 m and > 5 m fractions sampled at the chlorophyll maximum in the warm-core eddy had lower light requirements for photosynthesis than comparable surface samples from the same station. Results suggest that photoadaptation in natural populations ofSynechococcus spp. is accomplished primarily by changing photosynthetic unit number, occuring in conjuction with other accommodations in the efficiency of photosynthetic light reactions.     

Diel patterns of carbon incorporation into biochemical constituents of Synechococcus spp. and larger algae in the Northwest Atlantic Ocean

Diel patterns of ¹⁴C-bicarbonate incorporation in>5 m algal communities were compared with those in cyanobacterial populations of Synechococcus spp. (0.6 to 1.0 m), collected from the surface and/or chlorophyll maximum at three stations in the Northwest Atlantic Ocean (a neritic front; in Warm-Core Eddy 84-E; and Wilkinson's Basin) from 21 July to 8 August, 1984. Cell constituents were chemically separated into four fractions: lipids, low molecular weight (LMW) metabolites, polysaccharides/nucleic acids, and proteins. The in situ diel pattern of ¹⁴C assimilation was virtually the same for >5 m algal communities adapted to different environments. Protein synthesis appeared to continue at a reduced rate at night using energy derived from the catabolism of polysaccharides and the mobilization of LMW compounds. Synechococcus spp. populations exhibited inherent physiological differences in their in situ diel pattern of carbon fixation from that in>5 m algal communities taken from the same water mass. There was no nighttime protein-synthesis in Synechococcus spp. The relative proportion of ¹⁴C-protein remained constant over night, while that of ¹⁴C-polysaccharides/nucleic acids declined and that of labelled LMW metabolites increased. Daytime light-intensity manipulations did not alter the diel pattern of carbon fixation in any of the>5 m algal assemblages, while changes in the carbon metabolism of surface and shadeadapted Synechococcus spp. populations could be rapidly induced by altering the light intensity.Bigelow Laboratory Contribution No. 86004     

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.     

Growth,photosynthesis and carbon metabolism in the temperate marine diatom Skeletonema costatum adapted to low temperature and low photon-flux density

The temperate diatom Skeletonema costatum (Grev.) Cleve was grown in low temperature and/or low light conditions. The cultures were acclimatized for at least three months before experiments were begun. Our data indicate that the initial slope of the photosynthesis vs irradiance curve () is controlled predominantly by light history and the light-saturated photosynthesis (P _max) by temperature. The number of divisions per day decreased with decreasing light intensity, but was identical for cultures grown at 3° or 18°C. The metabolic pathways of inorganic carbon fixation were not fundamentally affected by low temperature or low light intensity, but both these factors increased labelling of C₃ compounds, synthesized by the Calvin-Benson cycle, and decreased that of phosphoenolpyruvate (PEP) and other metabolites. This indicates an enhancement of ribulose-1,5-bisphosphate (RuBP) carboxylase activity, which is the first step in the C₃ pathway (3-phosphoglycerate and sugar phosphate synthesis); this may optimize cell functions. At low temperatures, a seven-fold increase in RuBP carboxylase activity per cell was observed. S. costatum is able to adapt to low irradiance by increasing and decreasing I _k (the ratio of P _max:, light intensity at onset of light saturation), and to low temperature by increasing its cellular chlorophyll a and RuBP carboxylase content. However, in the latter case, adaptation is not optimal. This study revealed two main features: (1) there is evidence that RuBP carboxylase has a key function in adjustment to high rates of photosynthesis at suboptimal temperatures or irradiances; (2) adaptive mechanisms are dynamic processes and the role of the time scale in physiological adaptation should be considered.     

Diurnal patterns of size-fractioned primary productivity across a coastal front

In July 1985, diurnal patterns of photosynthesis and pigmentation were characterized for whole water (>0.4 m) and size-fractioned (>5 m and 0.4 to 5 m) communities from three light depths sampled across a coastal thermal front in the Southern California Bight. Samples were collected predawn and held for 20 h in deck incubators. Variations in chlorophyll a and accessory pigment-to-chlorophyll a ratios showed no obvious diurnal trends. Timing of peak photosynthetic potential (P _max) and its coincidence with variations in light-limited rates of photosynthesis (alpha), as well as diurnal amplitudes in P _max and alpha, often differed between size fractions sampled within the same community. The same was true for identical size fractions collected from different depths and stations transecting the front. Primary productivity was 20-fold greater on the cold water side, where >5 m diatoms dominated the mixed layer and accounted for 80% of daytime productivity. Diatoms collected from the top and bottom of the upper mixed layer displayed nearly identical diurnal patterns in P _max and alpha, with midday peaks exceeding predawn values by four-fold and two-fold respectively. Above the pycnocline, the 0.4 to 5 m fraction had lower assimilation rates than the >5 m fraction and smaller diurnal amplitudes in P _max and/or alpha, with daytime patterns often characterized by two peaks interspersed by a short period of photoinhibition. Within the front, the 0.4 to 5 m fraction accounted for two-thirds of plant biomass and >90% of primary production. Pigment analyses by high-performance liquid chromatography revealed enrichment in 19-hexanoyloxyfucoxanthin, indicative of enhanced numbers of prymnesiophtes. Photosynthetic activity in confined surface communities was susceptible to daytime photoinhibition, but subsurface communities exhibited midday P _max peaks that were three-to seven-fold predawn values. In the warm-water mass, both algal size fractions contributed equally to photosynthesis and chlorophyll a in surface waters, with the 0.4 to 5 m fraction becoming dominant at the base of the euphotic zone. At all depths, peak P _max of the 0.4 to 5 m fraction occurred before noon, while P _max of the >5 m fraction was clearly evident in the afternoon. Elevated chlorophyll b-, 19hexanoyloxyfucoxanthin- and zeaxanthin-to-chlorophyll a ratios indicated a mixture of algal groups, including chlorophytes, cyanobacteria and prymnesiophytes.     

Diel periodicity of photosynthesis in marine phytoplankton

Short-term changes in photosynthesis were documented for 17 of 24 marine phytoplankton species, representing a range of taxonomic groups. Periodicity in phytoplankton photosynthesis on light-dark cycles (diel periodicity) was widespread but not universal for the species studied. The centric diatoms Lauderia borealis, Ditylum brightwellii, Stephanopyxis turris, Coscinodiscus rex, Chaetoceros gracile, and Biddulphia mobiliensis had strong diel periodicity in photosynthetic capacity (P _max). Amplitudes of the daily variations ranged from 2.9 to >50, with maxima in the morning or near midday, and with minima during the dark period, and these variations were not dependent on changes in cell pigmentation. There was some evidence for sustained photosynthetic periodicity in constant conditions in several diatoms, and an endogenous rhythm may have been present. The photosynthesis-irradiance (P-I) relationship was time-dependent for representative marine diatoms, with both the initial slope () and the asymptote (P _max) of P-I curves exhibiting significant synchronous diel oscillations. Moreover, detailed studies of the amplitude and timing of photosynthetic periodicity for the diatoms L. borealis and D. brightwellii demonstrated large temporal variations in photosynthesis with morning maxima. These P-I oscillations are discussed with reference to models of primary production which use the relationship between photosynthesis and light as a component of predictive equations for phytoplankton growth in the sea.     

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     

Effects of nitrate on the diurnal vertical migration,carbon to nitrogen ratio,and the photosynthetic capacity of the dinoflagellate Gymnodinium splendens

A non-thecate dinoflagellate, Gymnodinium splendens, was studied in a 12 d laboratory experiment in 2.0x0.25 m containers in which light, temperature, and nutrients could be manipulated. Under a 12 h light: 12 h dark cycle, the dinoflagellates exhibited diurnal vertical migrations, swimming downward before the dark period began and upward before the end of the dark period. This vertical migration probably involved geotaxis and a diel rhythm, as well as light-mediated behavior. The vertical distribution of nitrate affected the behavior and physiology of the dinoflagellate. When nitrate was present throughout the container, the organisms resembled those in exponential batch culture both in C:N ratios and photosynthetic capacity (P_max); moreover, they migrated to the surface during the day. In contrast, when nitrate was depleted, C:N ratios increased, P_max decreased, and the organisms formed a subsurface layer at a depth corresponding to the light level at which photosynthesis saturated. When nitrate was present only at the bottom of the tank, C:N ratios of the population decreased until similar to those of nutrient-saturated cells and P_max increased; however, the dinoflagellates behaved the same as nutrient-depleted cells, forming a subsurface layer during the light period. Field measurements revealed a migratory subsurface chlorophyll maximum layer dominated by G. splendens. It was just above the nitracline during the day, and in the nitracline during the night, which concurs with our laboratory observations.     

Effects of altered photic regimes on diel patterns of species-specific photosynthesis

Diel patterns of photosynthesis were measured for two polar diatoms (Coscinodiscus sp. and Porosira pseudodenticulata) collected in September 1985 from McMurdo Sound, Antarctica, and four temperate dinoflagellates (Gonyaulax hyalina, Gymnodinium splendens, Dinophysis caudata, and Glenodinium sp.) collected in July 1985 and January 1986 from the Southern California Bight, California, USA. For phytoplankton incubated under three combinations of photoperiod and irradiance, distinct diel patterns of light-saturated (P _max) and light-limited (P _L ) photosynthesis were found for (i) different species isolated from the same environment, and (ii) polar diatoms and temperate dinoflagellates. The time of day when the maximum rate of P _max occurred was influenced by both irradiance and daylength for the polar diatoms but not by daylength for three out of four temperate dinoflagellates. The range of values of the ratio of maximum to minimum rates of photosynthesis (P _max:P _min) was similar for polar diatoms and temperate dinoflagellates. The results of this study suggest that changes in irradiance or photoperiod could influence species-specific patterns of photosynthesis in nature. As a consequence, in light-limited environments differential reproductive success could result from these diel patterns, and ultimately be reflected in temporal and spatial differences in community structure.     

The theoretical basis for estimating phytoplankton production and specific growth rate from chlorophyll, light and temperature data

Phytoplankton maximum specific growth rate, μ_max, and maximum photosynthetic quantum yield, Φ_max, can be related mathematically via the photosynthetic light curve, P(I). A model is presented in which maximum quantum yield defines the initial slope of the light curve and is assumed to be a known constant, while maximum specific growth rate defines the light-saturated region of the curve and is assumed to be a known function of temperature. The effect of introducing μ_max(T) into P(I, Φ_max) is to replace the unknown, temperature-dependent light saturation parameter with a term involving the ratio μ_max/Φ_max. The advantage of writing P(I) in terms of both μ_max and Φ_max is that those parameters are particularly well documented in the literature. Consequently, estimates of nutrient-unlimited phytoplankton growth and production rates can be based solely on the constants μ_max, Φ_max and k_c (light absorption per unit of chlorophyll) and the free variables light, temperature and chlorophyll concentration. Rate estimates appear to be accurate to within a factor of two for an extremely wide range of conditions.One particularly significant result of introducing μ_max into P(I, Φ_max) is that the carbon : chlorophyll ration, θ, appears explicitly. It is possible to derive an expression for optimum θ based on the assumption that adaptive changes in carbon/chlorophyll occur so as to maximize the specific growth rate for given conditions of light and temperature. Laboratory and field data are compiled from the literature to test the formulae presented here.     

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.     

Photosynthetic characteristics of marine phytoplankton from contrasting physical environments

Two suites of phytoplankton samples have been collected in consecutive years at various times over a day from selected depths within vertically mixed and stratified water columns in the western Irish Sea, in order to provide a range of possible light histories within the populations collected. Values for the maximum rate of ¹⁴C retention (P _max) and the initial slope of the ¹⁴C retention: light intensity curve () were obtained. Supra-thermocline samples from the stratified water exhibited higher P _max values than corresponding subthermocline samples. Higher values of were also generally associated with samples from the supra-thermocline zone of the stratified region. Differences in the depth distribution of P _max and in the mixed water were small, except in the presence of a shallow thermocline. In one suite of samples from the stratified water, a diurnal increase in the P _max values of the supra-thermocline samples was observed. P _max values obtained from the samples from the mixed water were interpreted in relation to the distribution obtained from the samples from the stratified zone. Data from both the contrasting sites visited for one sample suite demonstrated a two-phase relationship between the chlorophyll a concentration and both P _max and . The rates of ¹⁴C retention of the first suite of samples were estimated by two techniques. The average differences in the retention were greater in samples from the sub-as opposed to suprathermocline zone. No trends were apparent in the smaples from the mixed waters.     

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.     

Vertical habitat-partitioning by copepodites and adults of subtropical oceanic copepods

Vertical distributions of the abundant larger copepods, both adults and late copepodites, were observed day and night in the upper 500 m of the North Pacific central gyre in early November, 1971. Densities of the copepodites usually equalled or exceeded those of the adults. Copepod species with maximum densities at or above 100 m (Calanus spp., Nannocalanus minor f. major, Undinula darwini, and Euchaeta rimana) usually had no ontogenetic or diel migration. Neocalanus spp. and Haloptilus longicornis exhibited ontogenetic but not diel migrations. Nannocalanus minor f. minor, Aetideus acutus, Euchaeta media, Scolecithrix spp. and Pleuromamma spp., had both ontogenetic and diel migrations. Adults and copepodites of E. media and Pleuromamma spp. usually had their night modes at the same depth, but the daytime modes were at progressively deeper depths for progressively older stages. Daytime modes for adults and copepodites of A. acutus and Scolecithrix bradyi were at the same depth, but the nighttime modes were at shallower depths for progressively older stages. Night modes of all these migrators were usually in the mixed layer (75 m), where primary production rates were maximal. Congeners usually had similar migratory behavior, but competition probably has been a significant determinant of vertical distribution, since congeners, particularly sibling species, consistently had different depths of maximum occurrence during both day and night.     

Mesocosm studies of DCMU-enhanced fluorescence as a measure of phytoplankton photosynthesis

When measurements of in-vivo fluorescence are used to estimate photosynthesis in the field, the marked temporal and spatial variations in phytoplankton populations, and their nutrient and light histories, have produced varied results. Natural phytoplankton populations in large, flow-through mesocosms with different controlled nutrient and sewage sludge additions were sampled weekly from June to September 1984. Good correlations were observed between the increase in fluorescence upon the addition of DCMU (F) and both in-situ production and the parameters (P_m and ) of the photosynthesis-irradiance curve for these phytoplankton populations. Good correlations were also obtained between DCMU-enhanced fluorescence (F_DCMU) and chlorophyll a concentrations. The relationship between F and in-situ ¹⁴C production was consistent among mesocosms even in the face of major shifts from diatom-dominated to dinoflagellate-dominated populations. On the other hand, the F_DCMU:Chl a relationship was significantly different between mesocosms and related to species composition. It was concluded that F offers the possibility of rapidly and accurately indexing both in-situ production and the photosynthetic capacity of mixed phytoplankton populations.This study was supported by EPA cooperative agreement 810265-03 and NOAA grant NA-83-ABD-00008     

Photosynthetic characteristics of phycoerythrin-containing marine Synechococcus spp.

Marine Synechococcus spp. are sufficiently abundant to make a significant contribution to primary productivity in the ocean. They are characterized by containing high cellular levels of phycoerythrin which is highly fluorescent in vivo. We sought (Jan.–Apr., 1984) to determine the adaptive photosynthetic features of two clonal types of Synechococcus spp., and to provide a reliable physiological basis for interpreting remote sensing data in terms of the biomass and productivity of this group in natural assemblages. It was found that the two major clonal types optimize growth and photosynthesis at low photon flux densities by increasing the numbers of photosynthetic units per cell and by decreasing photosynthetic unit size. The cells of clone WH 7803 exhibited dramatic photoinhibition of photosynthesis and reduction in growth rate at high photon flux densities, accompanied by a large and significant increase in phycoerythrin fluorescence. Maximal photosynthesis of cells grown under 10–50 E m^-2 s^-1 was reduced by 20 to 30% when the cells were exposed to photon flux densities greater than 150 E m^-2 s^-1. However, steady-state levels of photosynthesis maintained for brief periods under these conditions were higher than those of cells grown continuously at high photon flux densities. No photoinhibition occurred in clone WH 8018 and rates of photosynthesis were greater than in WH 7803. Yields of in-vivo phycoerythrin fluorescence under all growth photon flux densities were lower in clone WH 8018 compared to clone WH 7803. Since significant inverse correlations were obtained between phycoerythrin fluorescence and P_max and for both clones grown in laboratory culture, it may be possible to provide a reliable means of assessing the physiological state, photosynthetic capacity and growth rate of Synechococcus spp. in natural assemblages by remote sensing of phycoerythrin fluorescence. Poor correlations between phycoerythrin fluorescene and pigment content indicate that phycoerythrin fluorescence may not accurately estimate Synechococcus spp. biomass based on pigment content alone.     

Diel relationships of microbial trophic groups and in situ dissolved carbohydrate dynamics in the caribbean sea

Dissolved total carbohydrate (TCHO), polysaccharide (PCHO), monosaccharide (MCHO) and organic carbon (DOC) were determined at 3-h intervals over 5 diel cycles in the mixed layer of the northwestern Caribbean Sea while following a drogued buoy. These data have been compared to populations of phototrophic (PNAN) and heterotrophic (HNAN) nanoplankton (2–20 m diameter) and heterotrophic bacteria (HBAC) (0.2–2.0 m diameter) estimated by epifluorescence counts, as well as to CO₂, phosphate, chlorophyll a and phaeopigment data determined simultaneously. Two different types of apparent diel dissolved carbohydrate (CHO) patterns were found. On 3 d when no sustained net CO₂ uptake was evident, TCHO and PCHO generally declined during the afternoon and early evening while MCHO tended to increase. On two other days when apparent sustained CO₂ uptake occurred during the day, there were large evening TCHO and PCHO peaks with constant or declining MCHO levels. These accumulations probably resulted from the release of recently produced PCHO from phototrophs. As was found earlier in the Sargasso Sea, PNAN populations were inversely related to PCHO concentrations. The sample to sample fluctuations of PNAN also were inversely related to the apparent rates of change of TCHO and PCHO, possibly due to an inverse relation between the rates of PNAN cell division and CHO excretion. Fluctuations in HBAC populations were inversely correlated with PCHO dynamics and directly related to MCHO variations, possibly due to extracellular hydrolysis of PCHO to MCHO during periods of rapid bacterial growth as well as to net heterotrophic PCHO uptake. A direct relationship between HNAN and TCHO fluctuations suggests the importance of HNAN excretion in the release of dissolved organics. The combined PNAN and HBAC fluctuations accounted for a more significant fraction of the variance in the apparent rates of change of PCHO than did any single population parameter indicating that intimate interactions between the microbial plankton groups are important in the in-situ regulation of CHO dynamics. Total system net TCHO release and uptake rates for 5 d averaged 56 and 53 g C l^-1 d^-1 respectively, assuming that the observed fluctuations resulted from temporal planktonic processes in homogeneous water masses. While the data contain indications that this was the case, this assumption is not definitive.     

Phytoplankton and pigment distributions in an anticyclonic slope water oceanic eddy (SWODDY) in the southern Bay of Biscay

An anticyclonic slope water oceanic eddy (SWODDY), named AE6, was sampled in the southern Bay of Biscay from 12 to 31 August 1998 to assess changes in the abundance and composition of phytoplankton assemblages related to the mesoscale feature. SWODDY AE6 showed characteristic biological signatures. A twofold increase in chlorophyll a concentration was found at the eddy centre relative to surrounding waters. Picoplankton cells accounted for a lower fraction of total chlorophyll a values at the eddy centre (44–50%) than outside the eddy (54–61%). Microscopic cell counts and HPLC pigment analysis showed that diatoms were almost entirely confined to the eddy centre, but both techniques yielded different results when studying other phytoplankton groups. Microscopic cell counts indicated that the spatial distribution of diatoms, dinoflagellates and unidentified flagellates was significantly influenced by SWODDY AE6, showing maximum abundance inside the mesoscale feature. HPLC pigment analysis provided more detailed information about the composition of pico–nanoplanktonic organisms. Pigment data processed by means of the CHEMTAX program showed "chlorophytes", "haptophytes" and "dinoflagellates II" (having haptophyte-like pigments and gyroxanthin-diester) as the more abundant "pigment classes" at the eddy centre, whereas dominance of "chlorophytes" and higher contribution of "cyanobacteria" (type Synechococcus) were estimated in the surrounding waters.Communicated by S.A. Poulet, Roscoff     

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.