Using a phytoplankton growth model to predict the fractionation of stable carbon isotopes

In the preceding paper in this issue, a phytoplankton growthmodel based on an analogy with chemical kinetics (the CR model)was re-derived, and a comparison made with the growth rate ofcultured phytoplankton assemblages extracted from temperatelakes. In this paper, further derivation of the CR model leadsto the same model of carbon isotope fractionation used by Rauet al. (Mar. Ecol. Prog. Ser., 133, 275–285, 1996). Boththe CR and Rau et al. models are compatible with the observationthat isotope fractionation during phytoplankton growth,     

The unsteady continuous culture of phosphate-limited Monochrysis lutheri droop: Experimental and theoretical analysis

A three-state variable model for phosphate-limited phytoplankton growth in a continuously lit continuous culture is proposed. In the model, the phosphate uptake rate per cell is a Michaelis-Mententype hyperbolic function of ambient nutrient concentration and the growth rate is a Droop-type hyperbolic function of cell quota. Steady-state and short-term uptake experiments with unialgal cultures of Monochrysis lutheri Droop, a marine chrysophyte, were used to calibrate the proposed model. For the long-term unsteady experiments, the model predicts well the culture's dynamic response in terms of cell density to steps down and up in influent concentration of limiting nutrient. For step changes in dilution rate, the model predicts well the culture's response to a step down but predicts poorly the culture's response to a step up. The long-term responses of the cultures to impulses in influent concentration show that the model fails to predict, even qualitatively, the behavior of the phytoplankton. Not unexpectedly, the model fails most dramatically in those experiments involving a rapid increase in cell quota, thereby demonstrating both the inherent flaws in the concept of the instantaneous growth rate as a function of instantaneous cell quota and the need for further dynamic characterization of phytoplankton behavior.     

Respiration and respiratory electron transport activity in marine phytoplankton: growth rate dependence and light enhancement

Respiratory electron transport system (ETS) activity and actualoxygen consumption rates were measured in batch cultures offour species of marine phytoplankton, in two different growthstages: exponential or log-phase (L) and stationary phase (S).The L cultures showed higher ETS activity and respiration ratesthan the S cultures of the same species. Among the L cultures,the higher respiration and ETS activity corresponded to thosehaving higher growth rates. The carbon-specific ETS activityand the carbon-specific respiration (h^–1) showed a cleardependence on growth rates. Samples subjected to short (10 min)exposures to high, oversaturating irradiances (1000 µEm^–2 s^–1) displayed enhanced ETS activity and respiration.The experiments show that, under the light regime at which thealgal cells grow, the respiratory ETS activity and actual oxygenconsumption in phytoplankton are strongly related to growthrate and that short, high irradiance exposures enhance boththe respiratory enzyme activity and their actual oxygen consumption.     

Modelling the effects on phytoplankton communities of changing mixed depth and background extinction coefficient on three contrasting lakes in the English Lake District

1. The process‐based phytoplankton community model, PROTECH, was used to model the response of algal biomass to a range of mixed layer depths and extinction coefficients for three contrasting lakes: Blelham Tarn (eutrophic), Bassenthwaite Lake (mesotrophic) and Ullswater (oligotrophic). 2. As expected, in most cases biomass and diversity decreased with decreasing light availability caused by increasing the mixed depth and background extinction coefficient. The communities were generally dominated by phytoplankton tolerant of low light. Further, more novel, factors were identified, however. 3. In Blelham Tarn in the second half of the year, biomass and diversity did not generally decline with deeper mixing and the community was dominated by nitrogen‐fixing phytoplankton because that nutrient was limiting to growth. 4. In Bassenthwaite Lake, changing mixed depth influenced the retention time so that, as the mixed depth declined, the flushing rate in the mixed layer increased to the point that only fast‐growing phytoplankton could dominate. 5. In the oligotrophic Ullswater, changing the mixed depth had a greater effect through nutrient supply rather than light availability. This effect was observed when the mixed layer was relatively shallow (<5.5 m) and the driver for this was that the inflowing nutrients were added to a smaller volume of water, thus increasing nutrient concentrations and algal growth. 6. Therefore, whilst changes in mixed depth generally affect the phytoplankton via commonly recognized factors (light availability, sedimentation rate), it also affected phytoplankton growth and community composition through other important factors such as retention time and nutrient supply.     

Allelopathic effect of the aquatic macrophyte, Stratiotes aloides, on natural phytoplankton

1. A survey of different Dutch Stratiotes stands showed that the density of phytoplankton (except cyanobacteria) was always higher outside S. aloides than between the rosettes of S. aloides. Analyses of water samples revealed that nutrient limitation was unlikely to have caused the lower phytoplankton biomass in the vicinity of S. aloides. 2. An in situ incubation experiment in the Danube Delta, Romania, indicated allelopathic activity against phytoplankton in S. aloides stands. The growth rate of natural phytoplankton populations exposed to water from S. aloides stands was significantly lower than that of populations that had not been in contact with S. aloides exudates. 3. A laboratory microcosm experiment showed a significantly lower phytoplankton biomass in treatments with S. aloides exudates. Nutrient concentrations and the light intensity were high enough that the lower phytoplankton biomass could not be explained by nutrient or light limitation.     

Phytoplankton growth in relation to network topology: time‐averaged catchment‐scale modelling in a large lowland river

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A model of physiological adaptation in unicellular algae

A simple growth model for unicellular algae is used to show that environmentally induced changes in cellular composition can be explained in terms of controlled adjustments acting to maximize the specific growth rate. The model is based on a division of cellular carbon into four distinct compartments: carbon associated with the photosynthetic apparatus, carbon associated with those components engaged in macromolecular synthesis, carbon associated with structural components and stored carbon. Flows of material between compartments, and between cells and their environment, are defined in terms of the environmental conditions and the distribution of carbon amongst compartments. Given that growth is balanced under a specific set of environmental conditions, there exists a unique, optimal allocation of carbon for which the rate of growth is maximal. Changes in this optimal allocation of material induced by changes in light intensity, nutrient availability or temperature are qualitatively similar to compositional changes observed in a wide variety of algal species. Empirical estimates for each of the model parameters are derived and used to show that reasonable quantitative agreement between observed and predicted behaviour is attainable. The model and parameter set are also used to illustrate the influence of cell size on growth rate. Under a given set of environmental conditions, the function relating cell size to growth rate has a single maximum. The size at which growth rate is maximal varies inversely with light intensity and directly with nutrient availability and temperature. Such behaviour is consistent with some empirical observations on the influence of environmental factors on the size distribution of natural phytoplankton communities.     

Seasonal succession of phytoplankton in an ice-free pond warmed by a thermal power plant

In a pond receiving warmed cooling waters from a thermal power plant, the physical and chemical properties of the water, phytoplankton, periphyton and zooplankton were monitored on a weekly sampling schedule. In winter the phytoplankton growth was limited by poor light conditions. In mid-February a rapid phytoplankton growth started, simultaneously with increasing light energy, high nutrient concentrations and small herbivorous zooplankton populations. The increase of phytoplankton biomass was stopped by lack of free nutrients and silica at the end of March. From May until August the phytoplankton standing crop was mainly regulated by herbivorous zooplankton. The autumnal maximum of phytoplankton occurred with decreasing zooplankton populations, increasing nutrient concentrations, a turbulence favourable for diatoms and high water temperature.     

Temporal and vertical dynamics of phytoplankton net growth in Castle Lake, California

The impact of nutrient additions, zooplankton grazing and light intensity on phytoplankton net growth with depth and season was studied with five microcosm experiments in meso-oligotrophic, subalpine Castle Lake, California, during the period of summer stratification in June-September 1994. The incubations (4 day) were performed at 5 m intervals from the surface to the bottom using natural phytoplankton and zooplankton assemblages, with enrichments of phosphorus and nitrogen. The phytoplankton community was only limited by nutrients in the upper 5 m (epilimnion), as indicated by change in chlorophyll concentration. Nutrient enrichments had the greatest effect on the phytoplankton net growth in June and July. High light inhibited the phytoplankton net growth at the surface. Low light intensities limited phytoplankton at 20 m and below, and at the end of the growing season already around 10-15 m. A deep chlorophyll maximum in the hypolimnion in June-August was not limited by either light or nutrients. The results showed variation in grazers' impact on phytoplankton. These results suggest the importance of nutrient limitation only in the epilimnion with light inhibition at the surface, light limitation in the hypolimnion, and varying impact of zooplankton grazing in influencing the development of the phytoplankton in Castle Lake.      

Growth-rate dependent feeding rates in Daphnia pulicaria and Brachionus rubens: adaptation to intermediate time-scale variations in food abundance

The hypothesis was tested that zooplankton can adapt to fluctuationsof their food resources at intermediate time scales in orderto maximize their energy input. The cladoceran Daphnia pulicariaand the rotifer Brachionus rubens were cultured at a range offixed growth rates and then offered radioactively labelled algaeat limiting and non-limiting concentrations to determine theirfunctional response. Rotifers and cladocerans showed a remarkablysimilar response. Animals growing at low growth rates had highermaximum filtering rates, i.e. were better adapted to low foodconcentrations than those growing at high rates. The growthrate did not affect the maximum ingestion rate in any of thespecies, thus indicating that adaptation to the prevailing foodconditions was based on the process of food collection. Bothspecies were heavier when cultured at high growth rates. Plottingthe weight-specific maximum filtering rate versus the growthrate resulted in a pronounced negative slope. The slope forthe weight-specific maximum ingestion rate was also negative.Hence filtering and ingestion rates seem to be determined bythe size of the zooplankton, not by their weight.     

Growth Rates of Marine Phytoplankton: Correlation with Light Absorption by Cell Chlorophyll a

To better predict plant production in the sea, it would be desirable to be able to calculate, from easily obtainable measurements at one sampling, the growth rate of the prevailing stock of phytoplankton. To this end growth rates, pigment composition, cell volume and cell surface area data were collected for several species of marine phytoplankton in logarithmic growth at 20–21°C and 0.07 cal/cm². min light intensity. Similar data for one species, Dunaliella tertiolecta, are given for several combinations of light intensity and temperature, and for another species, Ditylum brightwellii, grown in nitrogen deficiency. The problem of estimating growth rates of phytoplankton was divided into three parts: 1) variation of growth rate among diverse species and its relationship to light absorption by cell chlorophyll a: 2) variation in growth rate with light intensity; 3) variation in growth rate with temperature. An equation has been formulated for calculating growth rate which provides a more precise fit of the data than do equations for growth rate based upon cell surface/volume ratios or cell volume. The formulation is based upon light absorption by chlorophyll a. It allows for variations in the efficiency of utilization of light absorbed by chlorophyll a and the changes in chlorophyll a content resulting from light intensity and temperature differences. We do not attempt to predict variations in growth rate with photoperiod or spectral distribution, nor do we allow for light effects upon growth rate not mediated by photosynthesis, so the model is, at best, a rough approximation of reality.     

Interactions between temperature and light intensity on growth and photosynthesis of the cyanobacterium Oscillatoria agardhii

Oscillatoria agardhii was grown in turbidostat cultures undera 16/8 h light/dark cycle at various combinations of light intensityand temperature. Temperature was found to influence only themaximal growth rate; this relationship was linear over the temperaturerange studied. An equation was derived describing the growthrate (µ) as a continuous function of light intensity andtemperature. The light harvesting pigments chlorophyll a andC-phycocyanin increased in concentration when growth becamelight limited. The regulation patterns observed did not suggestany influence of temperature on their steady state concentrations.The initial slope of the P versus I curves (     

Modelling phytoplankton productivity in turbid waters with small euphotic to mixing depth ratios

A model which was developed and calibrated for predicting algalgrowth in mass cultures, was modified for natural systems. Inturbid systems the ratio of euphotic to aphotic depth is usuallysmall and mixing may exceed the compensation depth. In orderto compensate for the various light regimes to which the phytoplanktonwould be subjected, the losses due to dark respiration weremodified so that the effective light history would determinethe actual rate. The efficiency of light utilization also changesunder different light regimes and the model was modified totake these variations into account. Both these modificationsresulted in different production profiles being generated forthe same surface conditions, but with different mixing depths,where the phytoplankton become more efficient as the light regimedeteriorates (i.e. less respiration and greater light utilizationefficiency). A further consequence is that the ‘criticalmixing depth’ is {small tilde}2.5 times greater than thatwhich was previously accepted, being {small tilde}20 times theeuphotic depth. The model predicted productivities to within90% of observed rates. It was also suggested how the predictionscould be used to determine the extent of nutrient limitation.The predictions also have biomanipulatory consequences, as alterationsof the light regime through the addition of non-photosynthesizingmaterials, under certain conditions, may even result in a stimulationof phytoplankton productivity.     

HETEROGENEITY OF CHLOROPLAST GAPDH GENES IN THE DINOPHYCEAE

The kinetics of xanthophyll-cycle pigment switching and fluorescence quenching dynamics in the marine diatom, Phaeodactylum tricornutum were determined in the context of dynamic and static growth light. Cultures were grown in a modified photobioreactor capable of producing dynamic light fields which exhibited attenuation characteristics similar to that of water; these cultures were pre-acclimated to high and low, static and dynamic, growth-light regimes for at least three days, and then examined under high, static and dynamic light. Pigment pools varied markedly. The two static light cultures had pigment complements that were very similar to "traditional" high and low-light static cultures. The dynamic-light grown cultures had pigment complements, which were very similar to each other but different from the static-grown cultures. The maximum xanthophyll-cycle pigment de-epoxidation state attainable under saturating light was equal for all four treatments. Induction of fluorescence quenching was significantly faster in the static-grown cultures, while xanthophyll-cycle de-epoxidation rates did not show as much variation. Minimum irradiances for xanthophyll-cycle induction were correlated to average growth irradiance. Taken as a whole, the results from this work suggest that dynamic light-grown phytoplankton have a unique photosynthetic functionality that is different from static light-grown phytoplankton. The significance of these observations in the context of realistic light fields, and the photosynthetic response capabilities of algae grown under them will be discussed.     

PHOTOSYNTHESIS IN THE DEEP BLUE SEA: PHOTOPHYSIOLOGICAL RESPONSES OF PHAEODACTYLUM TRICORNUTUM TO DYNAMIC AND STATIC IRRADIANCES

The kinetics of xanthophyll-cycle pigment switching and fluorescence quenching dynamics in the marine diatom, Phaeodactylum tricornutum were determined in the context of dynamic and static growth light. Cultures were grown in a modified photobioreactor capable of producing dynamic light fields which exhibited attenuation characteristics similar to that of water; these cultures were pre-acclimated to high and low, static and dynamic, growth-light regimes for at least three days, and then examined under high, static and dynamic light. Pigment pools varied markedly. The two static light cultures had pigment complements that were very similar to “traditional” high and low-light static cultures. The dynamic-light grown cultures had pigment complements, which were very similar to each other but different from the static-grown cultures. The maximum xanthophyll-cycle pigment de-epoxidation state attainable under saturating light was equal for all four treatments. Induction of fluorescence quenching was significantly faster in the static-grown cultures, while xanthophyll-cycle de-epoxidation rates did not show as much variation. Minimum irradiances for xanthophyll-cycle induction were correlated to average growth irradiance. Taken as a whole, the results from this work suggest that dynamic light-grown phytoplankton have a unique photosynthetic functionality that is different from static light-grown phytoplankton. The significance of these observations in the context of realistic light fields, and the photosynthetic response capabilities of algae grown under them will be discussed.     

Dynamics and limitations of phytoplankton biomass along a gradient in Mwanza Gulf,southern Lake Victoria (Tanzania)

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The vertical distribution of phytoplankton in stratified water columns

What determines the vertical distribution of phytoplankton in different aquatic environments remains an open question. To address this question, we develop a model to explore how phytoplankton respond through growth and movement to opposing resource gradients and different mixing conditions. We assume stratification creates a well-mixed surface layer on top of a poorly mixed deep layer and nutrients are supplied from multiple depth-dependent sources. Intraspecific competition leads to a unique strategic equilibrium for phytoplankton, which allows us to classify the distinct vertical distributions that can exist. Biomass can occur as a benthic layer (BL), a deep chlorophyll maximum (DCM), or in the mixed layer (ML), or as a combination of BL+ML or DCM+ML. The ML biomass can be limited by nutrients, light, or both. We predict how the vertical distribution, relative resource limitation, and biomass of phytoplankton will change across environmental gradients. We parameterized our model to represent potentially light and phosphorus limited freshwater lakes, but the model is applicable to a broad range of vertically stratified systems. Increasing nutrient input from the sediments or to the mixed layer increases light limitation, shifts phytoplankton towards the surface, and increases total biomass. Increasing background light attenuation increases light limitation, shifts the phytoplankton towards the surface, and generally decreases total biomass. Increasing mixed layer depth increases, decreases, or has no effect on light limitation and total biomass. Our model is able to replicate the diverse vertical distributions observed in nature and explain what underlying mechanisms drive these distributions.     

Implications of seasonal mixing for phytoplankton production and bloom development

Based on a 1D model considering phytoplankton and nutrients in a vertical water column, we investigate the consequences of temporal and spatial variations in turbulent mixing for phytoplankton production and biomass. We show that in seasonally mixed systems, the processes controlling phytoplankton production and the sensitivity of phytoplankton abundance to ambient light, trophic state and mixed-layer depth differ substantially from those at steady state in systems with time-constant diffusivities. In seasonally mixed systems, the annually replenished nutrient pool in the euphotic zone is an important factor for phytoplankton production supporting bloom development, whereas without winter mixing, production mainly depends on the diffusive nutrient flux during stratified conditions. Seasonal changes in water column production are predominantly determined by seasonal changes in phytoplankton abundance, but also by seasonal changes in specific production resulting from the transport of nutrients, the exploitation of the nutrient pool and the increase in light shading associated with phytoplankton growth. The interplay between seasonal mixing and the vertical distribution of mixing intensities is a key factor determining the relative importance of the processes controlling phytoplankton production and the sensitivity of the size and timing of the annual maximum phytoplankton abundance to the abiotic conditions.