EFFECTS OF ULTRAVIOLET‐A RADIATION AND NUTRIENT AVAILABILITY ON THE CELLULAR COMPOSITION OF PHOTOPROTECTIVE COMPOUNDS IN GLENODINIUM FOLIACEUM (DINOPHYCEAE)1

The photoprotective response in the dinoflagellate Glenodinium foliaceum F. Stein exposed to ultraviolet‐A (UVA) radiation (320–400 nm; 1.7 W · m²) and the effect of nitrate and phosphate availability on that response have been studied. Parameters measured over a 14 d growth period in control (PAR) and experimental (PAR + UVA) cultures included cellular mycosporine‐like amino acids (MAAs), chls, carotenoids, and culture growth rates. Although there were no significant effects of UVA on growth rate, there was significant induction of MAA compounds (28 ± 2 pg · cell^?1) and a reduction in chl a (9.6 ± 0.1 pg · cell^?1) and fucoxanthin (4.4 ± 0.1 pg · cell^?1) compared to the control cultures (3 ± 1 pg · cell^?1, 13.3 ± 3.2 pg · cell^?1, and 7.4 ± 0.3 pg · cell^?1, respectively). In a second investigation, MAA concentrations in UVA‐exposed cultures were lower when nitrate was limited (P < 0.05) but were higher when phosphate was limiting. Nitrate limitation led to significant decreases (P < 0.05) in cellular concentration of chls (chl c₁, chl c₂, and chl a), but other pigments were not affected. Phosphate availability had no effect on final pigment concentrations. Results suggest that nutrient availability significantly affects cellular accumulation of photoprotective compounds in G. foliaceum exposed to UVA.     

LOW THERMAL LIMIT OF GROWTH RATE OF SYMBIODINIUM CALIFORNIUM (DINOPHYTA) IN CULTURE MAY RESTRICT THE SYMBIONT TO SOUTHERN POPULATIONS OF ITS HOST ANEMONES (ANTHOPLEURA SPP.; ANTHOZOA,CNIDARIA)1

Symbiodinium californium (#383, Banaszak et al. 1993 ) is one of two known dinoflagellate symbionts of the intertidal sea anemones Anthopleura elegantissima, A. xanthogrammica, and A. sola and occurs only in hosts at southern latitudes of the North Pacific. To investigate if temperature restricts the latitudinal distribution of S. californium, growth and photosynthesis at a range of temperatures (5°C–30°C) were determined for cultured symbionts. Mean specific growth rates were the highest between 15°C and 28°C (μ 0.21–0.26 · d^?1) and extremely low at 5, 10, and 30°C (0.02–0.03 · d^?1). Average doubling times ranged from 2.7 d (20°C) to 33 d (5, 10, and 30°C). Cells cultured at 10°C had the greatest cell volume (821 μm³) and the highest percentage of motile cells (64.5%). Growth and photosynthesis were uncoupled; light‐saturated maximum photosynthesis (P_max) increased from 2.9 pg C · cell^?1 · h^?1 at 20°C to 13.2 pg C · cell^?1 · h^?1 at 30°C, a 4.5‐fold increase. Less than 11% of daily photosynthetically fixed carbon was utilized for growth at 5, 10, and 30°C, indicating the potential for high carbon translocation at these temperatures. Low temperature effects on growth rate, and not on photosynthesis and cell morphology, may restrict the distribution of S. californium to southern populations of its host anemones.     

THE INABILITY OF THE TEXAS “BROWN TIDE” ALGA TO USE NITRATE AND THE ROLE OF NITROGEN IN THE INITIATION OF A PERSISTENT BLOOM OF THIS ORGANISM1

A planktonic alga similar in general morphology and pigments to Aureococcus anophagefferens Hargraves and Sieburth has caused persistent and ecologically damaging blooms along the south Texas coast. Experiments using 100 μM NO₃^?, NO₂^?, and NH₄⁺ demonstrated that the alga could not use NO₃^? for growth but could use NO₂^? and NH₄⁺. Doubling iron or trace metal concentrations did not permit growth on NO₃^?. Chemical composition data for cultures grown in excess NO₃^? or NH₄⁺, respectively, were as follows: N·cell^?1 (0.88 vs. 1.3 pg), C:N ratio (25:1 vs. 6.4:1), C:chlorophyll a (chl a) (560:1 vs. 44:1), and chl a·cell^?1 (0.033 vs. 0.16 pg). These data imply that cells supplied with NO₃^? were N-starved. Culture addition of 10 mM final concentration chlorate (a nitrate analog) did not affect the Texas isolate while NO₃^? utilizing A. anophagefferens was lysed, suggesting that the NO₃^? reductase of the Texas isolate is nonfunctional. Rates of primary productivity determined during a dense bloom indicated that light-saturated growth rates were ca. 0.45 d^?1, which is similar to maximum rates determined in laboratory experiments (0.58 d^?1± 0.16). However, chemical composition data were consistent with the growth rate of these cells being limited by N availability (C:N 28, C:chl a 176, chl a·cell^?1 0.019). Calculations based on a mass balance for nitrogen suggest that the bloom was triggered by an input of ca. 69 μM NH₄⁺ that resulted from an extensive die-off of benthos and fish.     

OLISTHODISCUS LUTEUS (CHRYSOPHYCEAE). III. UPTAKE AND UTILIZATION OF NITROGEN AND PHOSPHORUS1

Uptake and assimilation of nitrogen and phosphorus were studied in Olisthodiscus luteus Carter. A diel periodicity in nitrate reductase activity was observed in log and stationary phase cultures; there was a 10-fold difference in magnitude between maximum and minimum rates, but other cellular features such as chlorophyll a, carbon, nitrogen, C:N ratio (atoms) · cell^?1 were less variable. K_s values (~2 μM) for uptake of nitrate-N and ammonium-N were observed. Phosphorus assimilated · cell^?1· day^?1 varied with declining external phosphorus concentrations; growth rates <0.5 divisions · day^?1 were common at <0.5 μM PO_4-P. Phosphate uptake rates (K_s= 1.0–1.98 μM) varied with culture age and showed multiphasic kinetic features. Alkaline phosphatase activity was not detected. Comparisons of the nutrient dynamics of O. luteus to other phytoplankton species and the ecological implications as related to the phytoplankton community of Narragansett Bay (Rhode Island) are discussed.     

Nutrient uptake and alkaline phosphatase (ec 3:1:3:1) activity of emiliania huxleyi (PRYMNESIOPHYCEAE) during growth under n and p limitation in continuous cultures

Emiliania huxleyi (strain L) expressed an exceptional P assimilation capability. Under P limitation, the minimum cell P content was 2.6 fmol P·cell^?1, and cell N remained constant at all growth rates at 100 fmol N·cell^?1. Both, calcification of cells and the induction of the phosphate uptake system were inversely correlated with growth rate. The highest (cellular P based) maximum phosphate uptake rate (V_max^P) was 1400 times (i.e. 8.9 h^?1) higher than the actual uptake rate. The affinity of the P‐uptake system (dV/dS) was 19.8 L·μmol^?1·h^?1 at μ = 0.14 d^?1. This is the highest value ever reported for a phytoplankton species. V_max and dV/dS for phosphate uptake were 48% and 15% lower in the dark than in the light at the lowest growth rates. The half‐saturation constant for growth was 1.1 nM. The coefficient for luxury phosphate uptake (Q_maxt/Q_min) was 31. Under P limitation, E. huxleyi expressed two different types of alkaline phosphatase (APase) enzyme kinetics. One type was synthesized constitutively and possessed a V_max and half‐saturation constant of 43 fmol MFP·cell^?1·h^?1 and 1.9 μM, respectively. The other, inducible type of APase expressed its highest activity at the lowest growth rates, with a V_max and half‐saturation constant of 190 fmol MFP·cell^?1·h^?1 and 12.2 μM, respectively. Both APase systems were located in a lipid membrane close to the cell wall. Under N‐limiting growth conditions, the minimum N quotum was 43 fmol N·cell^?1. The highest value for the cell N‐specific maximum nitrate uptake rate (V_max^N) was 0.075 h^?1; for the affinity of nitrate uptake, 0.37 L·μmol^?1·h^?1. The uptake rate of nitrate in the dark was 70% lower than in the light. N‐limited cells were smaller than P‐limited cells and contained 50% less organic and inorganic carbon. In comparison with other algae, E. huxleyi is a poor competitor for nitrate under N limitation. As a consequence of its high affinity for inorganic phosphate, and the presence of two different types of APase in terms of kinetics, E. huxleyi is expected to perform well in P‐controlled ecosystems.     

DIFFERENT RESPONSE MECHANISMS OF TWO PLANKTONIC DESMID SPECIES (CHLOROPHYCEAE) TO A SINGLE SATURATING ADDITION OF PHOSPHATE

Two planktonic algal species, Staurastrum chaetoceras (Schr.) G. M. Smith and Cosmarium abbreviatum Rac. var. planctonicum W. et G. S. West, from trophically different alkaline lakes, were compared in their response to a single saturating addition of phosphate (P) in a P-limited growth situation. Storage abilities were determined using the luxury coefficient R = Q_max/Q₀. Maximum cellular P quotas differed, depending on whether cells were harvested during exponential growth at μ_max (Q_max, R being 26.7 and 9.1 for C. abbreviatum and S. chaetoceras, respectively) or harvested after a saturating pulse at P-limited growth conditions (Q′_max, R being 53.5 and 20.2 for C. abbreviatum and S. chaetoceras, respectively). At stringent P-limited conditions, maximum initial uptake rates were higher in S. chaetoceras than in C. abbreviatum (0.094 and 0.073 pmol P·cell^?1·h^?1, respectively), but long-term (net) uptake rates (over ～20 min) were higher in C. abbreviatum than in S. chaetoceras (0.048 and 0.019 pmol P·cell^?1·h^?1, respectively). Before growth resumed after the onset of a large P addition (150 μmol·L^?1), a lag phase was observed for both species. This period lasted 2–3 days for S. chaetoceras and 3–4 days for C. abbreviatum, corresponding with the time to reach Q^′_max. Subsequent growth rates (over ～10 days) were 0.010 h^?1 and 0.006 h^?1 for S. chaetoceras and C. abbreviatum, respectively, being only 20%–30% of maximum growth rates. In conclusion, S. chaetoceras, with a relatively high initial P-uptake rate, short lag phase, and high initial growth rate, is well adapted to a P pulse of short duration. Conversely, C. abbreviatum, with a high long-term uptake rate and high storage capacity, appears competitively superior when exposed to an infrequent but lasting pulse. These characteristics provide information about possible strategies of algal species to profit from temporarily high P concentrations.     

KINETICS OF SILICIC ACID UPTAKE AND RATES OF SILICA DISSOLUTION IN THE MARINE DIATOM THALASSIOSIRA PSEUDONANA1,2

Tracer techniques using the stable isotope ³⁰Si were used to measure rates of silicic acid uptake and silica dissolution in silicon replete and silicon depleted populations of 2 clones of the marine diatom Thalassiosira pseudonana Hasle & Heimdal. Uptake kinetics were describable using the Michaelis-Menten equation for enzyme kinetics, and no threshold concentration for uptake was evident. The maximum specific uptake rate of the estuarine clone 3H (0.062–0.092 · h^?1) was higher than that of the Sargasso Sea clone 13-1 (0.028–0.031 · h^?1), but half-saturation constants for uptake by the 2 clones were not measurably different (0.8–2.3 μM for 3H; 1.4–1.5 μM for 13-1). There was little or no light dependence of uptake in populations grown under optimal light conditions prior to the experiment. Exponentially growing populations released silicic acid to the medium by dissolution of cellular silica at rates ranging from 6.5 to 15% of the maximum uptake rate.     

INFLUENCE OF IRRADIANCE ON THE FATTY ACID COMPOSITION OF PHYTOPLANKTON1

Eight species of marine phytoplankton commonly used in aquaculture were grown under a range of photon flux densities (PEDs) and analyzed for their fatty acid (FA) composition. Fatty and composition changed considerably at different PFDs although no consistent correlation between the relative proportion of a single FA and μ or chl a · cell^?1 was apparent. Within an individual species the percentage of certain fatty acids covaried with PFDs, growth rate and/or chl a · cell^?1. The light conditions which produced the greatest proportion of the essential fatty acids was species specific. Eicosapentaenoic acid. 20:5ω3 increased from 6.1% to 15.5% of the total fatty acids of Chaetoceros simplex Ostenfield grown at PFDs which decreased from 225 μE · m^?2· s^?1 to 6 μE · m^?2· s^?1, respectively. Most species had their greatest proportion of 20: 5ω3 at low levels of irradiance. Conversely, docosahexaenoic acid, 22:6ω3, decreased from 9.7% to 3.6% of the total fatty acids in Pavlova lutheri Droop as PFD decreased. The percentage of 22:6ω3 generally decreased with decreasing irradiances. In all diatoms the percentage of 16:0 was significantly correlated with PFD, and in three of five diatoms, with growth rate (μ). Results suggest that fatty acid composition is a highly dynamic component of cellular physiology, which responds significantly to variation in PFD.     

SURVEY FOR KARLOTOXIN PRODUCTION IN 15 SPECIES OF GYMNODINIOID DINOFLAGELLATES (KARENIACEAE,DINOPHYTA)1

Toxin analysis of 15 species of Kareniaceae revealed the presence of karlotoxin, KmTx 2, in only a single species (Karlodinium veneficum) but with variable activity in strains from the Swan (Km^SwanTx 2‐1, 2.1 pg · cell^?1; and Km^SwanTx 2‐2, 0.53 pg · cell^?1), Huon (Km^HuonTx 2, 0.86 pg · cell^?1), and Derwent rivers (<0.001 pg · cell^?1) in Australia. A newly isolated Southern Ocean species, Karlodinium conicum, contained a novel poorly hemolytic karlotoxin analogue (Km_conicumTx, 2.8 pg · cell^?1). The hemolytic potency (HD50%) of the Australian karlotoxins were as follows: Km^SwanTx 2‐1 (65.9 ± 4.8 ng) and Km^SwanTx 2‐2 (63.4 ± 3.7 ng), Km^HuonTx 2 (343 ± 4.9 ng), and Km_conicumTx (>4,000 ng). Species from the closely related genera Takayama (T. helix, T. tasmanica, T. tuberculata), Karenia (K. asterichroma, K. brevis, K. mikimotoi, K. papilionacea, K. umbella), and Karlodinium (Ka. australe, Ka. antarcticum, Ka. ballantinum, Ka. corrugatum, Ka. decipiens) were all consistently negative for karlotoxin production. Brevetoxin (PbTx) was only detected in K. brevis, and hemolytic activity was only observed in Ka. veneficum strains.     

Effects of diurnal fluctuations in light intensity on the efficiency of growth of Skeletonema costatum (Grev.) Cleve (Bacillariophyceae) in a cyclostat

The effects of diurnal variations in light intensity on the biomass characteristics and the efficiency of daily growth of Skeletonema costatum (Grev.) Cleve were evaluated. The relative importance of changes in carbon specific rates of respiration and organic release to the efficiency of growth was determined. Light intensity was either constant at 130 μE · m^?2 · s^?1 during the light period or fluctuated throughout the light period from 500 to 10 μE · m^?2 · s^?1 at rates of either 1 or 12 cycles · day^?1. Total daily light was equivalent for all light regimes at 5.6 E · m^?2 · day^?1.Daily rates of growth remained comparable at ≈ 1 · day^?1 under constant and fluctuating light regimes. Cell size as daily mean carbon · cell^?1, nitrogen · cell^?1 and cellular volume was decreased under diurnally varying light whereas daily mean chlorophyll a · cell^?1 was unaffected.Rates of respiration, organic release and gross production were elevated several fold under diurnally varying light in comparison to constant light. Net growth efficiency decreased from 0.69 under constant light to values of 0.50 and 0.38 under 1 and 12 cycles · day^?1, respectively. Decreased efficiency of growth under diurnally fluctuating light resulted mostly from greater respiratory activity while organic release remained < 10% of gross production. Increased rates of gross production reflected enhancement in the efficiency of carbon fixation with fluctuating light.     

GENITIC VARIABILITY IN TOXIN POTENCIES AMONG SEVENTEEN CLONES OF GAMBIERDISCUS TOXICUS (DINOPHYCEAE)1

Seventeen clones of the ciguatera-causing dinoflagellate Gambierdiscus toxicus Adachi and Fukuyo were acclimated to the same environments over several months. Significant variance components were detected between non-acclimated and acclimated cultures for cell potencies, yields and reproduction rates. The resultant variance in acclimated potencies among clones was statistically significant (P < 0.0001), indicating that potency can be used for genetic comparisons. However, cell potency differences for a clone of G. toxicus in the acclimated vs. non-acclimated phases can exceed genetic differences between clones. This stresses the need for a rigorous acclimation process. Caribbean isolates of G. toxicus were inherently more toxic than isolates from other areas. One Caribbean clone yielded 55 × 10^?4 mu (mouse units)·cell^?1 whereas clones Bermuda, the Bahamas, and Florida ranged from only 1.8 × 10^?4 mu·cell^?1 to a maximum of 19.8 × 10^?4 mu·cell^?1. Toxicity decreased with increasing latitude (r =–0.819, P < 0.01), indicating that environmental differences probably influenced the potencies. A comparison of acclimated reproduction rates at four light intensities also indicated that genetic differences among clones existed. The resulting reproduction rate/light slopes overlapped, indicating that the clones may be adapted to specific light regimes.     

UTILIZATION OF DISSOLVED ORGANIC CARBON BY NATURAL POPULATIONS OF EPIBENTHIC SALT MARSH DIATOMS1

Natural populations of pennate diatoms collected from the mudflats of a Georgia salt marsh were cleansed by differential centrifugation and tested for their ability to assimilate and respire ¹⁴C-acetate, lactate and glucose at 1 μM concentrations. A correction was made for contaminating bacterial activity. Utilization rates were measured in the light and dark and in the presence of a second substrate. Mean incorporation rates were (in pmol ·μg chl a^?1· h^?1± SE): acetate, 117 ± 37; lactate, 53.7 ± 20.4; glucose 13.0 ± 5.8. Using a carbon/chlorophyll ratio of 53.2 and a generation time of 48 h we calculate these algae could obtain up to 1% of their carbon heterotrophically if several usable substrates were each available at 1 μM concentrations.     

PHYSIOLOGICAL INDICATORS OF NUTRIENT DEFICIENCY IN CLADOPHORA (CHLOROPHYTA) IN THE CLARK FORK OF THE COLUMBIA RIVER,MONTANA1

Cellular nutrient concentrations and nutrient uptake rates of Cladophora glomerata (L.) Kuetzing were determined during summer and fall in 1989–1990 at a site on the upper Clark Fork of the Columbia River, Montana. Both physiological tests indicated that Cladophora growth is likely to be limited by nitrogen during late summer-early fall. Maximum uptake rates of ammonia-N and nitrate-N were 5935–6991 and 507–984 μg · g DW^?1· h^?1, respectively, during July–October when dissolved inorganic nitrogen (DIN) concentrations in the river were less than 10 μg · L^?1. During November-December, when DIN was 72–376 μg · L^?1, maximum ammonia-N uptake was 1137–1633 μg · g DW^?1· h^?1 and maximum nitrate-N uptake was 0–196 μg · g DW^?1· h^?1. Cellular nitrogen during summer–early fall was 0.78–1.80% of Cladophora dry weight, frequently at or below 1.1%, a level suggested as a critical minimum N concentration for maximum growth. In contrast, cellular P was 0.18–0.36% of dry weight, 3–6 times the suggested critical P concentration of 0.06%. Molar ratios of cellular N:P (< 16:1) and DIN: SRP (< 4:1) during late summer-early fall also indicated potential N limitation. Cellular N and P from Cladophora collected from a second site influenced by a municipal wastewater discharge in 1990 displayed similar seasonal trends. At both sites, seasonal fluctuations in DIN were closely tracked by changes in cellular N, Cellular P, however, increased through the growing season despite declining levels of SRP in the river.     

EFFECT OF TEMPERATURE,LIGHT AND pH ON GROWTH,PHOTOSYNTHESIS AND RESPIRATION OF THE DINOFLAGELLATE PERIDINIUM CINCTUM FA. WESTII IN LABORATORY CULTURES1

Optimum light, temperature, and pH conditions for growth, photosynthetic, and respiratory activities of Peridinium cinctum fa. westii (Lemm.) Lef were investigated by using axenic clones in batch cultures. The results are discussed and compared with data from Lake Kinneret (Israel) where it produces heavy blooms in spring. Highest biomass development and growth rates occurred at ca. 23° C and ≥50 μE· m^?2·s¹ of fluorescent light with energy peaks at 440–575 and 665 nm. Photosynthetic oxygen release was more efficient in filtered light of blue (BG 12) and red (RG 2) than in green (VG 9) qualities. Photosynthetic oxygen production occurred at temperatures ranging from 5° to 32° C in white fluorescent light from 10 to 105 μE·m^?2·s^?1 with a gross maximum value of 1500 × 10^?12 g·cell^?1·h^?1 at the highest irradiance. The average respiration amounted to ca. 12% of the gross production and reached a maximum value of ca. 270·10^?12 g·cell^?1·h^?1 at 31° C. A comparison of photosynthetic and respiratory Q₁₀-values showed that in the upper temperature range the increase in gross production was only a third of the corresponding increase in respiration, although the gross production was at maximum. Short intermittent periods of dark (>7 min) before high light exposures from a halogen lamp greatly increased oxygen production. Depending on the physiological status of the alga, light saturation values were reached at 500–1000 μE·m^?2·s^?1 of halogen light with compensation points at 20–40 μE·m^?2·s^?1 and I_k-values at 100–200 μE·m^?2·s^?1. The corresponding values in fluorescent light in which it was cultured and adapted, were 25 to 75% lower indicating the ability of the alga to efficiently utilize varying light conditions, if the adaptation time is sufficient. Carbon fixation was most efficient at ca. pH 7, but the growth rates and biomass development were highest at pH 8.3.     

TOXIN PROFILE,PIGMENT COMPOSITION,AND LARGE SUBUNIT RDNA PHYLOGENETIC ANALYSIS OF AN ALEXANDRIUM MINUTUM (DINOPHYCEAE) STRAIN ISOLATED FROM THE FLEET LAGOON,UNITED KINGDOM1

Paralytic shellfish toxins, pigment composition, and large subunit (LSU) rDNA sequence were analyzed for a clonal culture of Alexandrium minutum Halim isolated in 2000 from the coastal Fleet Lagoon, Dorset, United Kingdom. The HPLC pigment analysis revealed the presence of chl a, peridinin, and diadinoxanthin as major pigments and chl c₁+c₂ and c₃, diatoxanthin, and β‐carotene as minor components. The toxins responsible for paralytic shellfish poisoning were analyzed by HPLC with postcolumn derivatization and fluorescence detection. The paralytic shellfish poisoning toxin profile of the Fleet Lagoon strain of A. minutum in exponential growth phase was dominated by gonyautoxin‐3 up to 54%, whereas gonyautoxin‐2 made up 10% and saxitoxin (STX) 36%. The average toxicity of the culture was 3.8 pg STX Eq·cell^?1, and total toxin content varied from 5.6 fmol·cell^?1 on day 1 to a maximum of 16.8 fmol·cell^?1 during the early stationary phase. Sequence analysis of the LSU rDNA revealed the strain to be closely related to several European strains of A. minutum and one isolated from Australian waters, although most of these do not produce STX. The shallow Fleet Lagoon may provide a favorable environment for A. minutum to bloom, and the presence of highly potent saxitoxins in this strain indicates potential for future shellfish contamination.     

CELL‐SPECIFIC EXTRACELLULAR PHOSPHATASE ACTIVITY OF DINOFLAGELLATE POPULATIONS IN ACIDIFIED MOUNTAIN LAKES1

High bulk extracellular phosphatase activity (PA) suggested severe phosphorus (P) deficiency in plankton of three acidified mountain lakes in the Bohemian Forest. Bioavailability of P substantially differed among the lakes due to differences in their P loading, as well as in concentrations of aluminum (Al) and its species, and was accompanied by species‐specific responses of phytoplankton. We combined the fluorescently labeled enzyme activity (FLEA) assay with image cytometry to measure cell‐specific PA in natural populations of three dinophyte species, occurring in all the lakes throughout May–September 2007. The mean cell‐specific PA varied among the lakes within one order of magnitude: 188–1,831 fmol · cell^?1 · h^?1 for Gymnodinium uberrimum (G. F. Allman) Kof. et Swezy, 21–150 fmol · cell^?1 · h^?1 for Gymnodinium sp., and 22–365 fmol · cell^?1 · h^?1 for Peridinium umbonatum F. Stein. To better compare cell‐specific PA among the species of different size, the values were normalized per unit of cell biovolume (amol · μm^?3 · h^?1) for further statistical analysis. A step‐forward selection identified concentrations of total and ionic Al together with pH as significant factors (P < 0.05, Monte Carlo permutation test), explaining cumulatively 57% of the total variability in cell‐specific PA. However, this cell‐specific PA showed an unexpected reverse trend compared to an overall gradient in P deficiency of the lake plankton. The autecological insight into dinophyte cell‐specific PA therefore suggested other factors, such as light availability, mixotrophy, and/or zooplankton grazing, causing further PA variations among the acidified lakes.     

PHOTOACCLIMATION OF PHOTOSYNTHESIS IRRADIANCE RESPONSE CURVES AND PHOTOSYNTHETIC PIGMENTS IN MICROALGAE AND CYANOBACTERIA1

The photosynthesis‐irradiance response (PE) curve, in which mass‐specific photosynthetic rates are plotted versus irradiance, is commonly used to characterize photoacclimation. The interpretation of PE curves depends critically on the currency in which mass is expressed. Normalizing the light‐limited rate to chl a yields the chl a‐specific initial slope (α^chl). This is proportional to the light absorption coefficient (a^chl), the proportionality factor being the photon efficiency of photosynthesis (φ_m). Thus, α^chl is the product of a^chl and φ_m. In microalgae α^chl typically shows little (<20%) phenotypic variability because declines of φ_m under conditions of high‐light stress are accompanied by increases of a^chl. The variation of α^chl among species is dominated by changes in a^chl due to differences in pigment complement and pigment packaging. In contrast to the microalgae, α^chl declines as irradiance increases in the cyanobacteria where phycobiliproteins dominate light absorption because of plasticity in the phycobiliprotein:chl a ratio. By definition, light‐saturated photosynthesis (P_m) is limited by a factor other than the rate of light absorption. Normalizing P_m to organic carbon concentration to obtain P_m^C allows a direct comparison with growth rates. Within species, P_m^C is independent of growth irradiance. Among species, P_m^C covaries with the resource‐saturated growth rate. The chl a:C ratio is a key physiological variable because the appropriate currencies for normalizing light‐limited and light‐saturated photosynthetic rates are, respectively, chl a and carbon. Typically, chl a:C is reduced to about 40% of its maximum value at an irradiance that supports 50% of the species‐specific maximum growth rate and light‐harvesting accessory pigments show similar or greater declines. In the steady state, this down‐regulation of pigment content prevents microalgae and cyanobacteria from maximizing photosynthetic rates throughout the light‐limited region for growth. The reason for down‐regulation of light harvesting, and therefore loss of potential photosynthetic gain at moderately limiting irradiances, is unknown. However, it is clear that maximizing the rate of photosynthetic carbon assimilation is not the only criterion governing photoacclimation.     

A CARBON BUDGET FOR THE AUTOTROPHIC CILIATE MESODINIUM RUBRUM1

Extensive blooms of the autotrophic ciliate Mesodinium rubrum (Lohmann) occurred in the Peru coastal upwelling region at 15°S latitude in March through May 1977 and contributed significantly to the organic productivity of the region. From observations made during the JOINT-II oceanographic expedition, a budget of the carbon flux of these unusual photosynthetic organisms can be constructed. The light dependent C fixation was determined with short (1 h) incubations because of the organisms' sensitivity to confinement and rapid nutrient exhaustion. Maximum photosynthesis occurred at 50% of incident light with a maximum rate of particulate C synthesis of 2187 mg C · m^?3· h^?1. The specific carbon uptake rates were also high with a maximum light saturated value of 16.8 mg C ·(mg chl a)^?1· h^?1. The rate of excretion of dissolved organic C at the productivity maximum ranged from 16.1 to 181.1 mg C · m^?3· h^?1. The range of percent excretion was 1.8–12.5% the total C fixed, similar to the range found in both motile and nonmotile phytoplankton assemblages. Respiration, determined by the decrease in particulate C in the dark, averaged 4.6% of the previously fixed photosynthetic C · h^?1. M. rubrum actively took up amino acids and naturally occurring dissolved organic carbon. The C budget for this ciliate indicates that the daily contribution to the particulate food chain is large, although not as great as is indicated by short incubations. The contribution of M. rubrum to the productivity and elemental fluxes of upwelling and coastal ecosystems has been seriously underestimated.     

Microphytobenthic species composition,pigment concentration,and primary production in sublittoral sediments of the Trondheimsfjord (Norway)1

In spring 2005, monthly sampling was carried out at a sublittoral site near Tautra Island. Microphytobenthic identification, abundance (ABU), and biomass (BIOM), were performed by microscopic analyses. Bacillariophyceae accounted for 67% of the total ABU, and phytoflagellates constituted 30%. The diatom floristic list consisted of 38 genera and 94 species. Intact light‐harvesting pigments chl a, chl c, and fucoxanthin and their derivatives were identified and quantified by HPLC. Photoprotective carotenoids were also observed (only as diadinoxanthin; no diatoxanthin was detected). Average fucoxanthin content was 4.57 ± 0.45 μg fucoxanthin · g sediment dry mass^?1, while the mean chl a concentration was 2.48 ± 0.15 μg · g^?1 dry mass. Both the high fucoxanthin:chl a ratio (considering nondegraded forms) and low amounts of photoprotective carotenoids indicated that the benthic microalgal community was adapted to low light. Microphytobenthic primary production was estimated in situ (MPPs, from 0.15 to 1.28 mg C · m^?2 · h^?1) and in the laboratory (MPPp, from 6.79 to 34.70 mg C · m^?2 · h^?1 under light saturation) as ¹⁴C assimilation; in April it was additionally estimated from O₂‐microelectrode studies (MPP_O2) along with the community respiration. MPP_O2 and the community respiration equaled 22.9 ± 7.0 and 7.4 ± 1.8 mg C · m^?2 · h^?1, respectively. A doubling of BIOM from April to June in parallel with a decreasing photosynthetic activity per unit chl a led us to suggest that the microphytobenthic community was sustained by heterotrophic metabolism during this period.     

FATTY ACIDS IN PHOTOTROPHIC AND MIXOTROPHIC GYRODINIUM GALATHE‐ANUM (DINOPHYCEAE)

Fatty acids were measured in G. galatheanum grown either phototrophically, or mixotrophically with Storeatula major (Cryptophyceae) as prey. G. galatheanum, like many photosynthetic dinoflagellates, contains high amounts of n‐3 long‐chain‐polyunsaturated fatty acids (LC‐PUFA) such as docosahexaenoic acid (DHA, 22:6n‐3) and the hemolytic toxic fatty acid 18:5n‐3. We hypothesize that a benefit of phagotrophy in G. galatheanum is the acquisition of precursor linolenic acid (18:3n‐3) that fuels LC‐PUFA synthesis. Phototrophs grew at 0.37 d^?1, while mixotrophs grew at 0.40 d^?1 with a feeding rate of 0.62 d^?1. Photosynthesis was lower in mixotrophs (3.7 pg C cell^?1 h^?1) than phototrophs (4.9 pg C cell^?1 h^?1). DHA levels were higher in mixotrophs [3.7 (+/? 0.11) pg cell^?1] than phototrophs [3.0 (+/? 0.16) pg cell^?1] and prey [0.4 (+/? 0.01) pg cell^?1]. 18:5n–3 levels [1.7 (+/? 0.03) pg cell^?1] were similar in phototrophs and mixotrophs. An intermediate in n‐3 LC‐PUFA synthesis, 20:4n‐3, accumulated in mixotrophs [0.6 (+/? 0.27) pg cell^?1] relative to phototrophs (not detected) and prey [0.03 (+/? 0.002) pg cell^?1]. Low ratios of linolenic acid to DHA in phototrophic G. galatheanum (0.14) relative to mixotrophic G. galatheanum (0.29) and prey (2.14) are consistent with substrate limitation of LC‐PUFA synthesis in phototrophs. Accumulation of 20:4n‐3 suggests incomplete conversion of linolenic acid to DHA, possibly due to conditions in batch culture. We conclude that precursors for n‐3 LC‐PUFA biosynthesis in G. galatheanum may be acquired through ingestion of S. major, and may partially control feeding/photosynthesis in mixotrophic populations.