The daily integral of photosynthesis by Planktothrix rubescens during summer stratification and autumnal mixing in Lake Zürich

It has been suggested that the populations of planktonic cyanobacteria that occupy the metalimnion of stratified lakes during the summer months may be aestivating between the main periods of growth during entrainment in the epilimnion in spring and summer. We determined the vertical distribution of the biomass and daily integral of photosynthesis of the population of Planktothrix ( Oscillatoria ) rubescens in Lake Zürich for 136 d from July to November 1995. The population showed an 80-fold increase during the stratified period but it only doubled over the subsequent period of entrainment. During the first eight days, part of the increase was attributed to recruitment of filaments floating up from greater depths but all of the subsequent production could be accounted for by photoautotrophic growth. On sunny days the biomass-specific photosynthesis of this population reached some of the highest values over the whole period despite its depth (>13 m). On very cloudy days, however, primary productivity was very low and on 4 days, when the mean depth of the population exceeded 15 m, there was no net production. Over the whole period of the study, the accumulated photosynthetic production exceeded the increase in biomass of the population by a factor of 9·5. Although much of this production occurred during the period of entrainment only a small proportion was translated into growth of the population. It is concluded that the growth that takes place in the period of stratification in the metalimnion is essential to subsequent production.     

Stratification by cyanobacteria in lakes: a dynamic buoyancy model indicates size limitations met by Planktothrix rubescens filaments

The ability of the Planktothrix rubescens to stratify in Lake Zürich is related to the size and shape of the cyanobacterial filaments. Detailed measurements made in the lake are used in a dynamic computer model of buoyancy regulation to investigate the vertical movements of filaments tracking the depth at which the irradiance would support neutral buoyancy. The movement of the filament lags behind the constantly changing target depth owing to (a) the time taken for the filament to respond to the irradiance by changing its density and (b) the time it takes to move by sinking down or floating up through the water column. The model simulates the stratification depth over a 5-month period of the summer from the continuous measurements of irradiance and weekly measurements of light attenuation and temperature, without any further adjustment over the period. Models using filaments of the size observed in Lake Zürich explain several details of the observed depth changes: smaller planktonic cyanobacteria (e.g. Limnothrix sp.) are unable to migrate fast enough and larger ones (e.g. Anabaena spp.) will overshoot and become entrained in the epilimnion. The model can be used to simulate recruitment of Planktothrix filaments from different depths after vernal stratification. Recruitment of filaments from depths down to 45 m will contribute to the metalimnetic population increase in early July.     

The uptake of amino acids by the cyanobacterium Planktothrix rubescens is stimulated by light at low irradiances

The rates of uptake of five amino acids--alanine, glutamate, glycine, leucine and serine--by axenic cultures of the cyanobacterium Planktothrix rubescens were measured over a range of irradiances using the (14)C-labelled amino acids at the nanomolar concentrations observed in Lake Zürich. The rates in the light exceeded the dark rates by as much as two- to ninefold. The light-affinity constants for stimulation were similar, indicating a similar process for each of the five amino acids. The E(k) (light saturation irradiance) for light stimulation was only 1 micromol m(-2) s(-1), less than the compensation point for photosynthesis and autotrophic growth, and much lower than the E(k) for either process. The E(k) for amino acid uptake was also less than the irradiance at which filaments obtain neutral buoyancy, which determines the depth at which they stratify and the irradiance they receive. This indicates that stimulation of amino acid uptake by light of low irradiances provides a mechanism for supplementing growth of filaments stratifying deep in the metalimnion, which, while able to grow at low irradiances, are often left with insufficient light to sustain them. Acetate uptake was also stimulated by light, but the kinetics differed.     

The annual cycle of growth rate and biomass change in Planktothrix spp. in Blelham Tarn, English Lake District

SUMMARY 1. The changes in the vertical distributions of red coloured Planktothrix rubescens and green P. agardhii filaments in Blelham Tarn, English Lake District, were related to vertical profiles of temperature and light attenuation and to continuous records of the surface irradiance and windspeed, from August 1999 to October 2000. 2. The potential growth rate of each organism was calculated from the irradiance and temperature at 0.5 m depths and hourly intervals throughout the year, using algorithms determined from growth rates in culture. The analyses indicated that there was sufficient irradiance for growth, integrated over the 24‐h cycle, at depths down to the metalimnion where the Planktothrix populations stratified in summer. The compensation depth for growth by P. rubescens reached a maximum of 9.3 m in spring and midsummer, and fell to a minimum of 1.6 m in midwinter; the corresponding values for P. agardhii were 7.9 and 0.5 m. 3. The mixed depth (z_m) exceeded the critical depth for growth (z_b) by P. rubescens (the condition preventing population increase) on only 3 days of the year; for P. agardhii, however, z_m exceeded z_b on 31 days, contributing to its faster decline. The stratified population of P. rubescens was the major cause of light attenuation during the summer of 2000, and resulted in competitive exclusion of P. agardhii. 4. The calculated growth rates integrated over the depth of the water column in Blelham Tarn equalled, or exceeded, the measured changes of the populations during periods when they were increasing, during summer and autumn. Close agreement between the two values was found for much of the year when allowance was made for dilution of the lake population by rainfall over the watershed. During periods of rapid decline, of P. agardhii in September 1999, P. rubescens in December 1999 and both in July–August 2000, additional losses (e.g. by chytrid parasitism and grazing) are invoked.     

Comparative effects of the quality and quantity of light and temperature on the growth of Planktothrix agardhii and P. rubescens1

The effects of temperature, light intensity, and quality on the growth of the cyanobacteria Planktothrix agardhii (Gomont) Anag. et Komárek and P. rubescens (Gomont) Anag. et Komárek were assessed in batch cultures. The relative competitiveness of the green‐pigmented P. agardhii and the red‐pigmented P. rubescens was evaluated in separate and mixed cultures, under different light intensities and qualities (green, red, and white), and at two different temperatures, chosen as representative of the natural conditions favoring the respective blooms of each species. In monocultures, the P. rubescens strain appeared to be particularly well adapted to low intensities of green light and displayed a strong photoinhibition under high irradiance levels. The P. agardhii strain appeared less specialized with regard to light quality and also less sensitive to photoinhibition at higher irradiances. In competition experiments, temperature (15°C vs. 25°C) was the most important parameter in determining relative fitness of the species and competitive success. At 15°C, P. rubescens appeared to be much more competitive than P. agardhii, while P. agardhii was more competitive at 25°C. Under low irradiance, however, the pigmentation of these strains was of primary importance in determining the outcomes of the competition experiments. On the basis of our experimental results and on field observations, we propose that the successful growth leading to the proliferation of these two differently pigmented strains may largely depend on the combined conditions of light and temperature. The two strains, being genetically close relatives, could therefore be considered as two ecotypes that are adapted to different light and temperature environments. Competition experiments showed that the combination of these parameters largely controls the success of one strain in comparison to the other.     

Photoheterotrophy and light-dependent uptake of organic and organic nitrogenous compounds by Planktothrix rubescens under low irradiance

1. Planktothrix rubescens is the dominant photoautotrophic organism in Lake Zürich, a prealpine, deep, mesotrophic freshwater lake with an oxic hypolimnion. Over long periods of the year, P. rubescens accumulates at the metalimnion and growth occurs in situ at irradiance near the photosynthesis compensation point. Experiments were conducted to evaluate the contribution of photoheterotrophy, heterotrophy and light‐dependent uptake of nitrogenous organic compounds to the carbon and nitrogen budget of this cyanobacterium under conditions of restricted availability of light quanta. 2. We used both purified natural populations of P. rubescens from the depth of 9 m and an axenic culture grown under low irradiance at 11 μmol m^?2 s^?1 on a light : dark cycle (10 : 14 h) to determine the uptake rates of various amino acids, urea, glucose, fructose, acetate and inorganic carbon. The components were added to artificial lake water in low amounts that simulated the naturally occurring potential concentrations. 3. The uptake rates of acetate and amino acids (glycine, serine, glutamate and aspartate) were strongly enhanced at low irradiance as compared with the dark. However, no difference was observed in the uptake of arginine, which was taken up at high rates under both treatments. The uptake rates of glucose, fructose and urea were very low under all conditions. Similar results were obtained for both axenic P. rubescens and for purified natural populations of P. rubescens that were separated from bacterioplankton and other phytoplankton. 4. Metalimnetic P. rubescens that was stratified at low irradiance for weeks exhibited much higher uptake rates than filaments that were entrained in the deepening surface mixed layer and experienced higher irradiance. The added organic compounds contributed up to 62% to the total carbon uptake of metalimnetic P. rubescens. On the basis of a molar C : N ratio of 4.9, the nitrogen uptake as organic compounds satisfied up to 84% of the nitrogen demand. 5. The experiments indicate that photoheterotrophy and light‐dependent uptake of nitrogenous organic compounds may contribute significantly to the carbon and nitrogen budget of filaments at low irradiance typical for growth of P. rubescens in the metalimnion and at the bottom of the surface mixed layer.     

Cyanobacterial bloom termination: the disappearance of Planktothrix rubescens from Lake Bourget (France) after restoration

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Effect of temperature on growth of the cyanobacterium Aphanizomenon flos-aquae in Lake Biwa and Lake Yogo

The water bloom‐forming cyanobacterium Aphanizomenon flos‐aquae Ralfs ex Bornet et Flahault (Nos‐tocales, Cyanophyceae) appeared in Lake Biwa and Lake Yogo in 1999 for the first time. The morphological characteristics were described using natural samples. In contrast to the other water bloom‐forming cyanobacteria such as Microcystis and Anabaena in Lake Biwa and Lake Yogo, the small summer population of A. flos‐aquae is apt to grow in winter, suggesting the low temperature preference or tolerance of this species. In order to clarify the effect of temperature on the growth, culture experiments were conducted using an axenic strain isolated from Lake Biwa. The strain could grow at above 8°C with an optimum temperature ranging from 23 to 29°C, and survived even at 5°C for at least 25days under low light conditions. Although these results confirmed the ability of the bloom formation during late autumn and winter, it is still unclear why the Aphanizomenon bloom occurred at temperatures of ca 10°C in December and not immediately after the disappearance of Microcystis and/or Anabaena bloom during autumn.     

Effects of temperature and pH on growth and photosynthesis of the thermophilic cyanobacterium Synechococcus lividus as measured by pulse-amplitude modulated fluorometry

In this study, the effects of five different temperatures and pH conditions on growth and photosynthetic performance of Synechococcus lividus Copeland from Taiwan were monitored in the field and the laboratory by using an underwater pulse‐amplitude modulated (Diving‐PAM) fluorometer. In the field, the optimal growth temperature of S. lividus was found to be 57°C. Such a finding was congruent with the growth rate in the laboratory culture, in which the optimal growth temperatures ranged from 45 to 60°C. In photosynthetic performance, the light‐saturated maximum relative electron transport rate (ETR_max) and the light‐limited slope (α^ETR) exhibited highest values at 50°C. At five different pH conditions, higher ETR_max and α^ETR were observed from pH 7 to 9. In addition, regression analysis demonstrated a significant positive relationship between the growth rate and the ETR_max values (R² = 0.9527), indicating that the growth of S. lividus was largely restricted to its photosynthetic performance. In conclusion, the photosynthetic performance and growth of the thermophilic cyanobacterium S. lividus were sensitive to fluctuations in temperature but not in pH. The present investigation offers a better understanding of the photosynthetic physiology.     

Plasticity in relative growth rate and its components following a change in irradiance

A total of 244 plants from two species, Lythrum salicaria and Epilobium glandulosum, were grown individually in hydroponic sand culture from seed for 36 d. Until day 27 all plants experienced an irradiance of 550 μmol m^?2 s^?1 PFD and on day 27 half of the plants were subjected to a neutral shade treatment in which irradiance was reduced to 100 μmol m^?2 s^?1 photon fluy density (PFD). Measures of relative growth rate, net assimilation rate, specific leaf area, biomass partitioning to leaves, roots, structural tissues (i.e. stems, petioles and inflorescences) and tissue density were obtained from intensive harvests three or four times per day. The shade treatment caused an immediate decrease in relative growth rate and net assimilation rate. Within hours the specific leaf area of the shaded plants increased and leaf tissue density decreased, thus partially offsetting the decrease in relative growth rate. Biomass partitioning was not affected.     

Variation in the components of relative growth rate in 10 Acacia species from contrasting environments

In this study we assessed the inherent relative growth rate (RGR) under controlled environment conditions of 10 contrasting Acacia species from semi-arid and mesic environments. For several of the species, compound pinnate leaves produced early in the seedling stage, were gradually replaced by phyllodes (expanded petioles that form simple lamina). Other species either did not form phyllodes, or only did so to a minor degree by the end of the study. Phyllode production was dominant in the four slow-growing Acacia species from semi-arid environments (A. aneura, A. colei, A. coriacea and A. tetragonophylla), with leaf production being exclusive or dominant in five (A. dealbata, A. implexa, A. mearnsii, A. melanoxylon and A. irrorata) of the six faster-growing species from mesic environments. The exception was A. saligna which was fast growing but did produce phyllodes. From a carbon economy perspective, slow growth in the semi-arid species was not associated with lower net assimilation rates or less plant mass allocated to foliage. Rather, the primary factor associated with their slow growth was a smaller foliage area per unit foliage mass. This was true for comparisons based on the mean over all harvests or at set plant masses. The production of phyllodes by the semi-arid species substantially reduced foliage area per unit foliage mass, as this was lower for phyllodes than leaves in all species. To assess the impact that phyllode production had on ontogenetic changes in RGR, we modelled the situation where only leaves were formed. This analysis showed that changing from leaves to phyllodes substantially reduced the RGR. There was little difference in plant nitrogen concentration or the ratio of foliage nitrogen to plant nitrogen between the species. This resulted in foliage nitrogen productivity (dry mass gain per unit foliage nitrogen and time) being directly proportional to foliage area per unit foliage mass between species. We concluded that a smaller foliage area per unit foliage mass and phyllode production are the primary factors associated with lower RGR in contrasting Acacia species.     

Carbohydrate dynamics and growth rate of natural phytoplankton populations

Diurnal changes in carbohydrate content of natural phytoplankton populations differed from those found for cyanobacteria grown in continuous cultures. The carbohydrate accumulation rate was not constant during the light period. Also in contrast to results obtained using continuous cultures the photosynthetic characteristics changed during the light period. A close correlation was observed between changes in carbohydrate accumulation rate and the efficiency of photosynthesis over 24 hours. Seasonal changes in carbohydrate consumption rate over the dark period were proportional to changes in growth rate.     

Kinetics of photoautotrophic growth of Marchantia polymorpha cells in suspension culture

Chlorophyllous, cultured cells of Marchantia polymorpha L. (HYA-2 cell line) grow actively under photoautotrophic (lithotrophic) conditions. The maximum specific growth rate (μ_cell) was 0.64 day⁻¹ and the doubling time was 1.08 days under optimum conditions (165 μmol m⁻² s⁻¹, 1% carbon dioxide enriched atmosphere, 25^°C). The photosynthetic activity was 1.30 μmol CO₂-fixed (10⁶ cells)⁻¹ h⁻¹ [66 μmol (mg chlorophyll)⁻¹ h⁻¹] in the exponential phase. The growth course has two distinct phases, an exponential and a linear one. The exponential phase is observed as long as the population density is sufficiently low (less than 7.9 × 10⁶ cells ml⁻¹), so that practically all individual cells directly receive the full incident light. The effect of light on the specific growth rate is a linear function of photon flux density. Linear growth occurs after the population density is so high that the incident light is almost completely absorbed by the cell suspension. The growth rate is a logarithmic function of photon flux density, in contrast to the specific growth rate, and saturates at high photon flux densities. The conditions of maximum growth, however, are not wellbalanced between cell mass production and cell division. Therefore, the maximum growth does not continue for a long time.     

Evaluation of the maximum specific growth rate of a yeast indicating non-linear growth trends in batch culture

The maximum specific growth rate (μ_max) of an ethanolic D-xylose-fermenting yeast, Pichia stipitis, showing non-linear growth trends in batch culture, was calculated using the rate equation μ₂ = (1/Δt) ln(x ₂/x ₁). The absolute error Δμ, affecting μ₂, was derived using an equation given by Borzani (1994). Based on the assumption of linearity of growth curves between two closest time points, the relation between the two rate formulae, μ₁ = (1/xˉ)dx _t /dt and μ₂ = (1/Δt) ln(x ₂/x ₁) was established. In a particular condition, when μ₁ = μ₂, an equation has been developed, the roots of which are the specific growth rates at different time points. This revised version was published online in July 2006 with corrections to the Cover Date.     

Hybridoma growth and antibody production as a function of cell density and specific growth rate in perfusion culture

Steady state metabolic parameters for hybridoma cell line H22 were determined over a wide range of cell densities and specific growth rates in a filtration based homogeneous perfusion reactor. Operating the reactor at perfusion rates of 0.75, 2.0, and 2.9 day(-1)(each at four different specific growth rates), viable cell densities as high as 2 x 10(7) cells/mL were obtained. For the cell line under investigation, the specific monoclonal antibody production rate was found to be a strong function of the viable cell density, increasing with increasing cell density. In contrast, most of the substrate consumption and product formation rates were strong functions of the specific growth rate. Substrate metabolism became more efficient at high cell densities and low specific growth rates. The Specific rates of metabolite formation and the apparent yields of lactate from glucose and ammonia from glutamine decreased at low specific growth rates and high cell densities. While the specific oxygen consumption rate was independent of the specific growth rate and cell density, ATP production was more oxidative at lower specific growth rate and higher cell density. These observed shifts are strong indications of the production potential of high-density perfusion culture. (c) 1995 John Wiley & Sons, Inc.