Une approche vers l'estimation de la production potentielle du phytoplancton par analyse des cinétiques d'induction de fluorescence

In vivo chlorophyll fluorescence is particularly interesting ot ecologists because of various concepts (biomass, productivity, physiological state) associated with it. Using a modified spectrophotofluorometer, we have studied the kinetics of fluorescence in unialgal cultures and in a natural population of marine phytoplankton. Our apparatus did not achieve satisfactory results with cell suspensions having a chlorophyll concentration less than 10 g l^-1. We have also tested a method for estimating kinetics of diluted cultures and marine phytoplankton using cells collected on glass-fibre filters. For unialgal cultures in the exponential growth phase, the method proved satisfactory, and results obtained from both cell suspensions and filters were in good agreement. However, for aged cultures (principally diatoms) and natural marine phytoplankton the method proved unsuitable. The kinetics of fluorescence induction vary according to taxonomic position of the cells, light intensity of the measuring excitation beam and productiveness of the culture medium. The importance of the kinetics of fluorescence induction for characterization of phytoplankton activity is discussed.     

Nutrient-limited toxin production and the dynamics of two phytoplankton in culture media: A mathematical model

In this paper we have proposed and analyzed a simple mathematical model consisting of four variables, viz., nutrient concentration, toxin producing phytoplankton (TPP), non-toxic phytoplankton (NTP), and toxin concentration. Limitation in the concentration of the extracellular nutrient has been incorporated as an environmental stress condition for the plankton population, and the liberation of toxic chemicals has been described by a monotonic function of extracellular nutrient. The model is analyzed and simulated to reproduce the experimental findings of Graneli and Johansson [Graneli, E., Johansson, N., 2003. Increase in the production of allelopathic Prymnesium parvum cells grown under N- or P-deficient conditions. Harmful Algae 2, 135–145]. The robustness of the numerical experiments are tested by a formal parameter sensitivity analysis. As the first theoretical model consistent with the experiment of Graneli and Johansson (2003), our results demonstrate that, when nutrient-deficient conditions are favorable for the TPP population to release toxic chemicals, the TPP species control the bloom of other phytoplankton species which are non-toxic. Consistent with the observations made by Graneli and Johansson (2003), our model overcomes the limitation of not incorporating the effect of nutrient-limited toxic production in several other models developed on plankton dynamics.     

Binding of manganese by Antarctic Phaeocystis pouchetii and the role of bacteria in its release

The growth of Antarctic Phaeocystis pouchetii and associated bacteria in culture, and the binding and release of manganese were investigated using cultures derived from three clonal isolates collected from Prydz Bay, Antarctica, in November and December 1982. The cultured strains accumulated manganese from the culture medium. The concentration of Mn²⁺ in the alga was up to 58 times that of the medium. The Mn²⁺ apparently binds to the mucilage secreted by the cells and gives the alga its characteristic brown colour. Oxidation or complexing of Mn²⁺ by P. pouchetii was also indicated. Bacterial growth on the mucilage is inhibited in rapidly growing cultures by acrylic acid produced in the mucilage. Once the growth of P. pouchetii and production of ccrylic acid slow, bacterial numbers increase, leading to the solubilization of the mucilage and the release of Mn²⁺. In cool-temperate, subpolar and polar seas where P. pouchetii may form dense blooms, the binding and subsequent release of manganese may have important consequences for competing phytoplankton.     

Changes in the acitivy of fucoxanthin-excited photosynthesis in the marine diatom Phaeodactylum tricornutum grown under different culture conditions

Variations in the photosynthetic activity under monochromatic light was studied in Phaeodactulum tricornutum grown under various culture conditions, with special reference to the composition of photosynthetic pigments. Photosynthetic activity, under light-limiting conditions, was reduced when the cells were grown under strong light. The reduction was more extensive in activity resulting from fucoxanthin-excitation than in that from chlorophyll a-excitation. The diminution in activity for fucoxanthin-excited photosynthesis did not correlate with variations in the pigment content. A similar diminution was observed with chlorophyll a fluorescence upon excitation of fucoxanthin. The change was accelerated by lowering the culture temperature, or limiting the supply of nitrogen source. The results were interpreted in terms of a nitrogen-deficient state for algal cells induced by strong light, low temperature or a limited supply of nitrogen. This leads to a modification of the physicochemical state of in vivo fucoxanthin, so that the excitation energy of fucoxanthin is less efficiently transferred to chlorophyll a. The significance of the phenomenon in the oceanic primary production is discussed.     

Continuous measurement of the DCMU-induced fluorescence response of natural phytoplankton populations

The effect of 3x10^-6 M DCMU [3-(3,4-dichlorophenyl)-1, 1-dimethylurea] on in vivo chlorophyll a fluorescence was observed in nearshore waters of the Southern California Bight. We compared fluorescence readings in the presence and absence of this inhibitor using parallel flow-through fluorometers. The increase in fluorescence induced by DCMU is expressed as the FRI (fluorescence response index). Theory and laboratory studies on batch cultures of phytoplankton suggest that the FRI is correlated with photosynthetic efficiency and/or physiological state, but other studies have produced results in apparent conflict with this interpretation. Although sufficient information does not exist to justify the use of fluorescence response as a precise physiological indicator in the field, we suggest that very low FRI values are a manifestation of photosynthetic debility in a sample. Vertical profiles showed a wide range of the fluorescence response index. At a station close to shore, low FRI values were observed well below the 1% light level, but the fluorescence response of the phytoplankton throughout the euphotic zone was similar to that of growing cultures. Farther offshore, the FRI was depressed near the surface, but increased in the enhanced nutrient conditions of the lower euphotic zone. The patterns observed were strong, and consistent with hypotheses which relate low values of the FRI to diminished photosynthetic capacity.     

Interspecific differences in the photosynthetic carbon metabolism of marine phytoplankton

Six species of marine phytoplankton of different sizes and taxonomic categories were grown in microcosms under identical experimental conditions; the species cultured were: Pavlova lutheri (Prymnesiophyceae), Dunaliella tertiolecta (Chlorophyceae), Phaeodactylum tricornutum (Baciollariophyceae), Eutreptiella sp. (Euglenophyceae), Alexandrium tamarense (Dinophyceae), and Phaeocystis pouchetii (Prymnesiophyceae). The photosynthetic carbon metabolism of these phytoplankton was studied throughout the exponential and lag phases of growth after nutrient depletion. The relative incorporation of carbon into protein was positively correlated with phytoplankton growth, while carbon assimilation into low molecular weight metabolites (LMWM) and storage products, i.e., lipid and polysaccharides, generally increased under nutrient-limiting conditions. Clear taxonspecific differences were observed in the proportions of carbon incorporated into cell constituents. A significant linear relationship was consistently found between the relative synthesis of protein to LMWM, and both the production normalised to chlorophyll (P:B) and the phytoplankton growth rate. However, ANCOVA revealed significant, interspecific differences in these relationships.     

Stimulation and inhibition of phytoplankton growth by low molecular weight hydrocarbons

Experiments on 4 phylogenetically different phytoplankton exposed in culture to a range of concentrations of benzene, toluene and xylene showed a variety of growth responses for marine microalgae. The degree of influence of these aromatic hydrocarbons, all components of fuel oils and crude oils, varied with concentration, compound and species. Stimulation of growth in Dunaliella tertiolecta resulted from low μg/l concentrations of all three compounds, Skeletonema costatum showed no growth enhancement, while Cricosphaera carterae and Amphidinium carterae were intermediate in their reactions. Closed culture vessels were found to be necessary to retain these volatile hydrocarbons. Many of the previous laboratory studies on oil using standard methods — cotton plugs, screw caps or beakers — have overlooked the important influence of the volatile fraction. The species-specific stimulation of low concentrations was further shown in experiments with mixtures of No. 2 fuel oil. The volatile fraction was most biologically reactive, being the source of growth enhancement at low levels and a major growth inhibitor at high concentrations. Thus, a significant environmental effect of oil on marine primary production could be the growth stimulation of particular species by low molecular weight aromatic compounds resulting in an alteration of the natural phytoplankton community structure and its trophic relationships.     

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     

Experimental studies on the relationship between natural copper complexes and their toxicity to phytoplankton

Toxicity of copper to phytoplankton depends on the physicochemical form of the metal. Organic substances liberated into the culture medium by the phytoplankton species Cricosphaera elongata are able to detoxify and complex copper. The molecular weight range of these organic substances has been determined by ultrafiltration.     

Correlation between oxygen utilization and electron transport activity in marine phytoplankton

Measurement of a respiratory control ratio is deemed very useful in determination of energy balance in respiration. We have defined and computed a control ratio as the ratio of the rate of oxygen consumption in vivo to the calculated maximum rate of oxygen consumption in vitro as measured by enzymatic electron transport rates. In order for a control ratio to serve a useful purpose in calculating in situ oxygen consumption in a water parcel, the ratio must be fairly constant. Our results indicated that this is precisely so in healthy cultures of the 9 species of phytoplankton studied. The data presented in this communication further point towards the potential for making reasonable estimates of phytoplankton oxygen consumption in an ocean water parcel by directly utilizing in vitro electron transport activity measurements.     

Fluorescence properties of natural phytoplankton populations

The cellular fluorescence of chlorophyll a in natural phytoplankton was measured during vertical profiling in marine coastal waters. The ratio of in situ fluorescence to chlorophyll a concentration, which was considered as an index of cellular fluorescence, varied over a wide range, with large changes occurring both within the water column and between profiling sites. The variations were caused in part by an inhibition in the fluorescence of cells exposed to intense sunlight. The inhibition, which occurred at irradiances exceeding 0.15 langley (ly)/min, led to diel fluctuations in the fluorescence of those phytoplankton near the sea surface. The remaining variations were independent of changes in temperature, but were unexplained. Both light-dependent and light-independent variations in cellular fluorescence will affect the accuracy of the continuous, fluorometric measurement of in vivo chlorophyll.     

Phytoplankton photosynthetic activity and growth rates in the NW Adriatic Sea

Taxonomic composition, biomass, primary production and growth rates of the phytoplankton community were studied in two stations in the NW Adriatic Sea on a seasonal basis, in areas characterized by differing hydrological and trophic conditions. The main differences between the two stations were quantitative rather than qualitative, most phytoplankton species being common to both stations. The effects of differing nutrient concentrations and plume spreading were evident. Biomass and primary production rates were significantly higher in the coastal station (S1), and the phytoplankton distribution in the water column was markedly stratified in S1 and more even in the offshore station (S3). However, chlorophyll a specific production, potential growth rate and production efficiencies were very similar in both stations, even when phosphorus concentrations were limiting. A discrepancy between potential and actual growth rate was observed: as a feature common to both stations, comparisons between potential and actual growth rates revealed that little carbon produced by phytoplankton accumulated as algal biomass; therefore, very high loss rates were estimated.     

Persistent diel rhythms in the chlorophyll fluorescence of marine phytoplankton species

The diel periodicities of in-vivo chlorophyll fluorescence and DCMU-enhanced chlorophyll fluorescence of 47 marine phytoplankton species were examined for 2 d in a 14 h L:10 h D light: dark cycle and then in continous light for another 2 to 3 d. Almost all phytoplankton species exhibit a more rapid increase in in-vivo fluorescence and DCMU-enhanced fluorescence during the light phase than during the dark phase. About one-half of the species examined exhibited persistent diel rhythms in continous light, indicating the operation of a biological clock. No phylogenetic or habitat related trends as to which species exhibited persistent rhythms were apparent. Of the phyla of eukaryotic phytoplankton adequately examined, none lacked biological clocks. Contrary to past hypotheses, some phytoplankton species maintain a persistent diel rhythm in a constant environment while reproducing at a rate greater than one division per day.     

Depth-dependent changes in chlorophyll fluorescence number at a Sargasso Sea station

Chlorophyll a-specific in vivo fluorescence exhibited depth-dependent changes in a Sargasso Sea phytoplankton community, decreasing from a maximum value at the surface to a minimum at 90m, and then increasing again below 90 m. This distribution pattern was not explained by irradiance conditions, diurnal variability, senescence in the deep population, or changes in light-absorption efficiency of chlorophyll a. However, a significant positive correlation was found between mean phytoplankton cell size and fluorescence number in the upper euphotic zone, where nutrient concentrations were low. We hypothesize that the direct cause for this observed correlation was nutrient limitation. In this picoplankton-dominated community, packaging effect was minimal. Under nutrient-limiting conditions, as mean cell size increases photosynthetic efficiency decreases and therefore fluorescence number increases. In the lower euphotic zone where nutrients were not limiting, changes in fluorescence number exhibited weak size-dependence and appeared to be related to species compositional changes.     

Extensive cross-disciplinary analysis of biological and chemical control of Calanus finmarchicus reproduction during an aldehyde forming diatom bloom in mesocosms

Egg and faecal pellet production and egg hatching success of the calanoid copepod Calanus finmarchicus were monitored over a period of 14?days (14?C28 April, 2008) while fed water from 4 differently treated mesocosms and ambient water. Two of the mesocosms used were inoculated with the polyunsaturated aldehyde (PUA)-producing diatom Skeletonema marinoi, while 2 received only nutrient additions with or without silica. The mesocosms developed blooms of S. marinoi, mixed diatoms or the haptophyte Phaeocystis pouchetii, respectively. Faecal pellet production of C. finmarchicus increased with increasing food availability. Egg production increased with time in all mesocosms to a maximum single female production of 232 eggs female^?1?day^?1 (average of 90 eggs female^?1?day^?1) and followed the development of ciliates and P. pouchetii, but was not affected by the observed high (up to 15?nmol?L^?1) PUA production potential of the phytoplankton. The hatching success of the eggs produced on the mesocosm diets was high (78?C96%) and was not affected by either aldehydes in the maternal diet or exposure to the dissolved aldehydes in the water.     

Total organic sulfur and dimethylsulfoniopropionate in marine phytoplankton: intracellular variations

Cellular levels of particulate organic sulfur (POS) and dimethylsulfoniopropionate (DMSP) were determined in cultures of five species of marine phytoplankton, Amphidinium carterae, Prorocentrum minimum, Emiliania huxleyii, Dunaliella tertiolecta and Skeletonema costatum. The first three are known producers of DMSP while the latter two produce little or non-detectable amounts of DMSP. In those species which produced significant amounts of DMSP, intracellular levels of POS and DMSP varied for the dinoflagellates (A. carterae and P. minimum) while they remained fairly constant for the prymnesiophyte (E. huxleyii) over the growth cycle (until late stationary phase), and DMSP accounted for the majority of the POS (50 to 100%). In species with low levels of DMSP, intracellular POS and DMSP decreased or remained constant over the growth cycle, and DMSP accounted for a much lower percentage of the POS (0 to 40%). Species with high DMSP concentrations had higher POS levels per unit cell volume as well, and DMSP accounted for substantially higher percentages of the particulate organic carbon (POC). Molar N:S ratios suggest a non-protein origin for much of the sulfur in the species producing DMSP. During late stationary phase, an increasing percentage of the DMSP became extracellular in the dinoflagellate and diatom (S. costatum) cultures, suggesting leakage. In a bacterized algal culture, measured quantities were considerably less than in an axenic counterpart, suggesting consumption.     

Production primaire au niveau de la thermocline en zone néritique de Méditerranée Nord-Occidentale

Vertical profiles of physical, chemical and phytoplanktonic parameters are described, at the level of the thermocline, in the area of Banyuls-sur-Mer, France. The results show that the thermocline divides two masses of water: (1) Mediterranean surface water with low nutrient concentrations and a salinity below 38.00 ‰; (2) deep, nutrient-rich upwelled water (N?NO₃ >3 μat-g·l^-1, P?PO₄>0.3 μat-g·l^-1, >38.30 ‰ S), which comes from the upper limit of the Mediterranean intermediate water, usually located at the 200 m level. Consequently, conditions are suitable for high production rates at the bottom of the thermocline, where Chl a is above 0.5 mg·m^-3; dominant species are Nitzschia delicatissima and N. pungens. A diagram is presented explaining the different effects of the pycnoclines on primary production: eutrophication at the pycnocline levels is the result of passive accumulation of phytoplankton and organic matter during sedimentation, and/or of reduced diffusion of nutrients from deep waters towards the surface.     

Simulating the temporal pattern of waste production in farmed gilthead seabream (Sparus aurata), European seabass (Dicentrarchus labrax) and Atlantic bluefin tuna (Thunnus thynnus)

Most fish farming waste output models provide gross waste rates as a function of stocked or produced biomass for a year or total culture cycle, but without contemplating the temporality of the discharges. This work aims to ascertain the temporal pattern of waste loads by coupling available growth and waste production models and developing simulation under real production rearing conditions, considering the overlapping of batches and management of stocks for three widely cultured species in the Mediterranean Sea: gilthead seabream (Sparus aurata), European seabass (Dicentrarchus labrax) and Atlantic bluefin tuna (Thunnus thynnus). For a similar annual biomass production, the simulations showed that waste output and temporal dumping patterns differ between the three species as a result of the disparities in growth velocity, nutrient digestibility, maintenance metabolic budget and husbandry. The simulations allowed the temporal patterns including the periods of maximum discharge and the dissolved and particulate nitrogen and phosphorus content in the wastes released to be determined, both of which were seen to be species-specific.     

In situ grazing rate of the copepod population in the western subarctic North Pacific during spring

Time series sampling with a multi-layer plankton sampler was carried out in the western subarctic North Pacific during spring 1991. Neocalanus cristatus, N. flemingeri and Eucalanus bungii dominated and accounted for 88.5% of the copepod population in volume. Neocalanus spp. were distributed in the upper mixed layer, while E. bungii was mainly distributed between 120 and 300 m throughout the day and night. In contrast, Metridia pacifica, Pleuromamma scutullata and Gaetanus simplex showed clear diel vertical migration. Grazing activities were estimated simultaneously by gut fluorescence. Nocturnal grazing was observed for diel migrating species. Neocalanus spp. did not have a diel feeding rhythm and had relatively low gut fluorescence. E. bungii was considered to be dormant during the observation period. The estimated grazing rate of the copepod population on phytoplankton was 1.4 to 2.0% of the primary production while the metabolic requirement was 8.3 to 12.4% of the primary production. These facts suggest that the copepod population was unimportant as primary consumers and that microzooplankton plays a much more important role in sustaining low standing stock of phytoplankton and a high nutrient concentration in the western subarctic Pacific Ocean.     

Chlorophyll a fluorescence in marine centric diatoms: Responses of chloroplasts to light and nutrient stress

Observations at sea of large variations in the cellular fluorescence of phytoplankton prompted a study of the fluorescence responses in marine diatoms to light and nutrient stress. When older cultures of Lauderia borealis were exposed to intense light, the in vivo fluorescence of chlorophyll a declined within the first 2 min of exposure. This initial response to light stress appeared to be correlated with a contraction of the chloroplasts. Continued exposure led to a second decline in fluorescence, which required 30 to 60 min for completion. A movement of chloroplasts to the valvar ends of the cell caused this secondary response. Both the contraction and intracellular movement of chloroplasts appeared to be related to both photoinhibition of photosynthesis and diel fluctuations in cellular fluorescence. An investigation of continuous cultures of Cyclotella nana showed that in vivo chlorophyll a fluoresced more strongly in nitrogen-starved cells than in enriched ones. Photoinhibition of cellular fluorescence also increased with the cell's state of nitrogen deficiency.