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 Burgundy-blood phenomenon: a model of buoyancy change explains autumnal waterblooms by Planktothrix rubescens in Lake Zürich

Buoyancy changes of the cyanobacterium Planktothrix rubescens- the Burgundy-blood alga - were modelled from its buoyancy response to light and irradiance changes in Lake Zürich during autumnal mixing. The daily insolation received by filaments at fixed depths and circulating to different depths was calculated from the measured light attenuation and surface irradiance. The active mixing depth, za5, was determined from the vertical turbulent diffusion coefficient, Kz, calculated from the wind speed, heat flux and temperature gradients. The fixed depth resulting in filament buoyancy, zn, decreased from 13 to 2 m between August and December 1998; the critical depth for buoyancy, zq, to which filaments must be circulated to become buoyant, decreased from >60 m in the summer to <10 m in winter. When za5 first exceeded zn, in September, P. rubescens was mixed into the epilimnion. In October, zq > za5: circulating filaments would have lost buoyancy in the high insolation. Often in November and December, after deeper mixing and lower insolation, za5 > zq: filaments would have become buoyant but would have floated to the lake surface (the Burgundy-blood phenomenon) only under subsequent calm conditions, when Kz was low. The model explains the Burgundy-blood phenomenon in deeper lakes; waterblooms near shallow leeward shores arise from populations floating up in deeper regions of the lake.     

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.     

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.     

The trophic state of Lake Ladoga as indicated by late summer phytoplankton

As a part of the joint Russian-Finnish evaluation of human impact on Lake Ladoga, we studied the phytoplankton of the lake in order to find biological indicators for eutrophication. A second aim of the investigation was intercalibration of sampling and phytoplankton counting techniques between the Russian and Finnish laboratories. Phytoplankton samples were collected from 27 sampling stations in the lake and from the rivers Volkhov and Neva in 9–13 August 1993. In surface water samples the phytoplankton fresh weight biomass varied in the range 218–3575 mg m^–33. Highest biomass values were encountered in Sortavala Bay, and lowest ones in the western central part of the lake. Phytoplankton species composition varied considerably in the lake; blue-green and green algae predominated near-shore areas and Cryptophyceae in the offshore stations. Canonical correspondance analysis revealed close grouping of eutrophy indicating communities, dominated mainly by greens and blue-greens, in the most nutrient-rich parts of Lake Ladoga, the Volkhov and Svir Bays. Samples from the vicinity of the inflows of Vuoksi and Burnaya Rivers and off Pitkaranta formed a separate group, dominated by diatoms, most of which were typical to mesotrophic or eutrophy lakes. As judged by phytoplankton biomass values and chlorophyll a concentrations, Lake Ladoga may generally be classified as mesotrophic. Eutrophicated areas are found in the northern archipelago of the lake and in the areas influenced by large rivers.     

Phytoplankton succession in the euphotic zone of Lake Corlo (Northern Italy) during summer stratification

Abstract

Lake Corlo is a dammed reservoir for hydro-electrical purposes in Northern Italy. Weekly samplings were made from May to August 1985 in the euphotic layer. Two major categories, flagellates and diatoms, alternatively dominated the algal community. Flagellates prevailed with stratified conditions. The strong influence of hydraulic conditions on phytoplakton succession is emphasized.     

Thermal stratification, nutrient dynamics, and phytoplankton productivity during the onset of spring phytoplankton growth in Lake Baikal, Russia

Lake Baikal, Russian Siberia, was sampled in July 1990 during the period of spring mixing and initiation of thermal stratification. Vertical profiles of temperature, dissolved nutrients (nitrate and soluble reactive phosphorus), phytoplankton biomass, and primary productivity were determined in an eleven-station transect encompassing the entire 636 km length of the lake. Pronounced horizontal variability in hydrodynamic conditions was observed, with the southern region of the lake being strongly thermally stratified while the middle and north basins were largely isothermal through July. The extent of depletion of surface water nutrients, and the magnitude of phytoplankton biomass and productivity, were found to be strongly correlated with the degree of thermal stratification. Horizontal differences likely reflected the contribution of two important factors: variation in the timing of ice-out in different parts of the lake (driving large-scale patterns of thermal stratification and other limnological properties) and localized effects of river inflows that may contribute to the preliminary stabilization of the water column in the face of intense turbulent spring mixing (driving meso-scale patterns). Examination of the relationships between surface water inorganic N and P depletion suggested that during the spring and early summer, phytoplankton growth in unstratified portions of the lake was largely unconstrained by nutrient supplies. As summer progressed, the importance of co-limitation by both N and P became more apparent. Uptake and regeneration rates, measured directly using the stable isotope ¹⁵N, revealed that phytoplankton in stratified portions of the lake relied primarily on NH₄ as their N source. Rates of NH₄ regeneration were in approximate equilibrium with uptake; both processes were dominated by organisms <2 µm. This pattern is similar to that observed for oligotrophic marine systems. Our study underscores the importance of hydrodynamic conditions in influencing patterns of biological productivity and nutrient dynamics that occur in Lake Baikal during its brief growing season.     

The daily pattern of nitrogen uptake by phytoplankton in dynamic mixed layer environments

The transport and assimilation of the various forms of biologically available nitrogen by phytoplankton, and the subsequent biosynthesis of N-containing macromolecules, have the potential to respond in different ways during the daily growth cycle. This review examines five types of effect that may influence the daily pattern of nitrogen uptake and metabolism: light versus dark (the day/night cycle); changes in irradiance during the day (including the diurnal rise and fall in photon fluence rates); circadian rhythms (endogenous patterns of variation which may continue in the absence of external environmental forcing); periodic variations in exogenous nitrogen supply; and the 24-hour dynamics of stratification and mixing. The hydrodynamic effects operate through a variety of direct and indirect controls, and can substantially modify the diel rhythmicity of phytoplankton growth.     

The influence of the diel climatic cycle on the depth-time distribution of phytoplankton and photosynthesis in a shallow equatorial lake (Lake Baringo,Kenya)

Lake Baringo is a shallow equatorial lake. This paper reports a diel study of the depth-time distribution of phytoplankton and photosynthesis at one location in Lake Baringo on 10 March 1989. The water column shows a pattern of diurnal stratification probably accentuated by the high turbidity of the water and therefore rapid attenuation of solar energy. This stratified pattern breaks down at night due to atmospheric cooling and the regular onset of winds in the early evening. The phytoplankton is dominated byMicrocystis aeruginosa with some associated epiphytes. It concentrates in the narrow euphotic zone during the diurnal period of stratification due to buoyancy of theMicrocystis; evening breakdown of the thermocline results in the phytoplankton being mixed throughout the water column. A series of measurements of photosynthesis throughout the diurnal period gives an areal rate of 3.8 g O₂ m⁻² d⁻¹. The relationship between this value and the level of fish exploitation in Lake Baringo is discussed. The diel cycle in Lake Baringo is interpreted as dominating over any seasonal limnological cycle in the lake.     

The Physico-Chemical and Planktonic Response of an Algicide — Treated Shallow Mountain Lake in Arizona

The physico-chemical and planktonic response and recovery of a shallow mountain lake in Arizona was closely monitored for a 2 1/2 year period following treatment with an algicide. The influential role blue-green algal blooms exert on seasonal and annual lake dynamics was most obvious once nuisance populations were temporarily removed from the system. Differential sensitivity between algal species was observed. Cryptomonad flagellates were least sensitive to algicide treatment and were dominant during periods when dissolved organic materials appeared to increase. The effect of rotenone application on blue-green bloom species is discussed.     

The effects of diel changes in circulation and mixing on the longitudinal distribution of phytoplankton in a canyon‐shaped Mediterranean reservoir

1. The near‐surface distribution of phytoplankton cells along the thalweg of a canyon‐shaped reservoir (El Gergal, southern Spain) during two surveys is described and interpreted as the result of time‐varying large‐scale circulation patterns, vertical mixing processes and the physiological capacity of algal cells to regulate its position in the water column. 2. Vertical gradients of chlorophyll‐a concentration developed in the water column during the day but disappeared at night, as a result of the shoaling and deepening of the diurnal mixed layer (dml). The changes in the depth of the dml are largely controlled in El Gergal by convectively driven mixing processes. The longitudinal circulation changes, in turn, as a result of weak and diurnal land‐sea breezes. The distribution of algal cells was patchy at all times but did not change during any of the surveys. 3. An expression is proposed to estimate time scales for the development of horizontal patchiness T_P based on simple concepts of transport. It is shown that T_P is in the order of a week, indicating that horizontal patchiness does not respond immediately to hourly changes in the controlling factors. The magnitude of T_P, though, depends on how the vertical distribution of chlorophyll‐a and longitudinal currents change on subdiurnal time scales. In particular, T_P is sensitive to the lag existing between the momentum and heat fluxes through the free surface, driving circulation and vertical mixing.     

The relationship between carbon dioxide fixation and chlorophyll a fluorescence during induction of photosynthesis in maize leaves at different temperatures and carbon dioxide concentrations

The rate of CO₂ fixation (F_c) and 680 nm chlorophyll fluorescence emission (F680) were measured simultaneously during induction of photosynthesis in Zea mays L. leaves under varying experimental conditions in order to assess the validity of fluorescence as an indicator of in vivo photosynthetic carbon assimilation. Z. mays leaves showed typical Kautsky fluorescence induction curves consisting of a fast rise in emission (O to P) followed by a slow quenching via a major transient (S-M) to a steady-state (T). After an initial lag, net CO₂ assimilation commenced at a point corresponding to the onset of the S-M transient on the F680 induction curve. Subsequently, F_c and F680 always arrived at a steady-state simultaneously. Decreasing the dark-adaption period increased the rate of induction of both parameters. Alteration of leaf temperature produced anti-parallel changes in induction characteristics of F_c and F680. Reducing the CO₂ level to below that required for saturation of photosynthesis also produced anti-parallel changes during induction, however, at CO₂ concentrations tenfold greater than the atmospheric level the rate of F680 quenching from P to T was appreciably reduced without a similar change in the induction of F_c. Removal of CO₂ at steady-state produced only a small increase in F680 and a correspondingly small decrease in F680 occurred when CO₂ was re-introduced. The complex relationship between chlorophyll fluorescence and carbon assimilation in vivo is discussed and the applicability of fluorescence as an indicator of carbon assimilation is considered.Abbreviations F_c rate of CO₂ fixation - F680 fluorescence emission at 680 nm