Diurnal changes in buoyancy and vertical distribution in populations of Microcystisin two shallow lakes

Changes in the buoyancy of Microcystis populations were followedover 24 h periods in two shallow well-mixed lakes, Lake Vinkeveen(area 0 6 km²) and Lake IJsselmeer (1190 km²), in the NetherlandsThe Microcystis colonies collected from the surface layers inboth lakes showed a buoyancy decrease during the day and anincrease at night The buoyant colonies, and especially the faster-movinglarge ones, became concentrated by flotation into the surfacemixed layers As a result the mean position of the cyanobacterialpopulation became located nearer the surface than that of othernon-buoyant phytoplankton, such as Scencdesmus. The cyanobacteriawould, therefore, have received a higher average irradianceThe Microcystis in these shallow lakes had weaker gas vesiclesthan those found previously in deeper lakes but it was demonstratedthat the loss of buoyancy, which occurred at high irradiances,resulted from an increase in carbohydrate ballast rather thanthrough turgoi-driven gas vesicle collapse     

RECRUITMENT OF BENTHIC MICROCYSTIS (CYANOPHYCEAE) TO THE WATER COLUMN: INTERNAL BUOYANCY CHANGES OR RESUSPENSION?1

In some lakes, large amounts of the potentially toxic cyanobacterium Microcystis overwinter in the sediment. This overwintering population might inoculate the water column in spring and promote the development of dense surface blooms of Microcystis during summer. In the Dutch Lake Volkerak, we found photochemically active Microcystis colonies in the sediment throughout the year. The most vital colonies originated from shallow sediments within the euphotic zone. We investigated whether recruitment of Microcystis colonies from the sediment to the water column was an active process, through production of gas vesicles or respiration of carbohydrate ballast. We calculated net buoyancy, as an indication of relative density, using the amounts and densities of the major cell constituents (carbohydrates, proteins, and gas vesicles). Carbohydrate content of benthic Microcystis cells was very low throughout the year. Buoyancy changes of benthic Microcystis were mostly a result of changes in gas vesicle volume. Before the summer bloom, net buoyancy and the amount of buoyant colonies in the sediment did not change. Therefore, recruitment of Microcystis from the sediment does not seem to be an active process regulated by internal buoyancy changes. Instead, our observations indicate that attachment of sediment particles to colonies plays an important part in the buoyancy state of benthic colonies. Therefore, we suggest that recruitment of Microcystis is more likely a passive process resulting from resuspension by wind‐induced mixing or bioturbation. Consequently, shallow areas of the lake probably play a more important role in recruitment of benthic Microcystis than deep areas.     

Artificial mixing prevents nuisance blooms of the cyanobacterium Microcystis in Lake Nieuwe Meer, the Netherlands

1. Artificial mixing in the hypertrophic Lake Nieuwe Meer was successful in preventing blooms of the cyanobacterium Microcystis. During the 2 years of artificial, deep mixing the number of colonies of Microcystis per litre and also per m² was lower than in the two preceding control years. Hardly any nuisance scums of Microcystis occurred in the lake.
2. The phytoplankton shifted from a cyanobacteria-dominated community in summer to a mixed community of flagellates, green algae and diatoms. Reduced sedimentation losses in the mixed lake, probably in combination with a lower pH, favoured non-buoyant algae, while the entrainment of cyanobacteria in the turbulent flow nullified their advantage of buoyancy.
3. The chlorophyll concentrations were much lower in the mixed lake, but the euphotic depth did not show clear differences between the years. The chlorophyll content integrated through depth (m^–2) increased in the artificially mixed lake.
4. The deep lake normally stratified in summer, but artificial mixing of the lake in 1993 resulted in a homogeneous temperature and oxygen distribution with depth. In spring 1994, the mixing was applied intermittently with a reduction of 75% of the energy costs, while the mixing was still sufficient to prevent stratification.
5. Determination of the buoyancy state of the colonies on a sunny and calm day showed that the buoyancy loss was low close to the bubble plumes, and high at some distance from these plumes. This suggests that Microcystis could escape the mixing at some distance from the plumes, and could synthesize more carbohydrates during its stay in the upper illuminated layer of the lake than the deep mixed colonies close to the bubble plumes. Determination of the buoyancy state appeared to be a good and simple method to investigate the extent of entrainment of colonies in the turbulent flow.     

Artificial mixing prevents nuisance blooms of the cyanobacterium Microcystis in Lake Nieuwe Meer, the Netherlands

1. Artificial mixing in the hypertrophic Lake Nieuwe Meer was successful in preventing blooms of the cyanobacterium Microcystis. During the 2 years of artificial, deep mixing the number of colonies of Microcystis per litre and also per m² was lower than in the two preceding control years. Hardly any nuisance scums of Microcystis occurred in the lake.
2. The phytoplankton shifted from a cyanobacteria-dominated community in summer to a mixed community of flagellates, green algae and diatoms. Reduced sedimentation losses in the mixed lake, probably in combination with a lower pH, favoured non-buoyant algae, while the entrainment of cyanobacteria in the turbulent flow nullified their advantage of buoyancy.
3. The chlorophyll concentrations were much lower in the mixed lake, but the euphotic depth did not show clear differences between the years. The chlorophyll content integrated through depth (m^–2) increased in the artificially mixed lake.
4. The deep lake normally stratified in summer, but artificial mixing of the lake in 1993 resulted in a homogeneous temperature and oxygen distribution with depth. In spring 1994, the mixing was applied intermittently with a reduction of 75% of the energy costs, while the mixing was still sufficient to prevent stratification.
5. Determination of the buoyancy state of the colonies on a sunny and calm day showed that the buoyancy loss was low close to the bubble plumes, and high at some distance from these plumes. This suggests that Microcystis could escape the mixing at some distance from the plumes, and could synthesize more carbohydrates during its stay in the upper illuminated layer of the lake than the deep mixed colonies close to the bubble plumes. Determination of the buoyancy state appeared to be a good and simple method to investigate the extent of entrainment of colonies in the turbulent flow.     

RECRUITMENT OF MICROCYSTIS (CYANOPHYCEAE) FROM LAKE SEDIMENTS: THE IMPORTANCE OF LITTORAL INOCULA1

Recruitment of Microcystis from sediments to the water column was investigated in shallow (1–2 m) and deep (6–7 m) areas of Lake Limmaren, central Sweden. Recruitment traps attached to the bottom were sampled on a weekly basis throughout the summer season ( June–September). A comparison between the two sites showed that the recruitment from the shallow bay was significantly higher over the entire season for all three Microcystis species present in the lake. Maximum rates of recruitment were found in August, when 2.3 × 10⁵ colonies m ^{? 2}·day ^{? 1} 1 Received 18 April 2002. Accepted 29 October 2002. left the sediments of the shallow area. Calculated over the entire summer, Microcystis colonies corresponding to 50% of the initial abundance in the surface sediments were recruited in the shallow bay, whereas recruitment from the deep area was only 8% of the sediment colonies. From these results we conclude that shallow areas, which to a large extent have been overlooked in studies of recruitment of phytoplankton, may be crucial to the dynamics of these organisms by playing an important role as inoculation sites for pelagic populations.     

Seasonal variations of Microcystis populations in sediments of Lake Biwa, Japan

Seasonal variations of colony numbers of Microcystis aeruginosa(Kütz.) Kütz. and M. wesenbergii(Komárek) Komárek in N. V. Kondrat. in sediments of Lake Biwa were investigated over a period of 1 year. At two stations located in the shallow South Basin of Lake Biwa (ca. 4 m water depth), the colony number of Microcystisfluctuated seasonally. The number had a tendency to gradually decrease from winter to early summer, while it increased through mid-summer and autumn. Since the Microcystispopulation in sediment was rather small, intensive growth and accumulation in the water column should be important for the formation of Microcystisblooms in Lake Biwa. Microcystiscolonies in the sediment samples after June were observed to be floating in a counting chamber under a microscope. The observation suggests that the recruitment of Microcystis colonies into the water column mostly occurs in early summer. The number of Microcystiscolonies in the deep North Basin of Lake Biwa (70 – 90 m water depth) was larger than in the South Basin. Because the seasonal variation of colony numbers was not observed in the North Basin, and Microcystiscells do not have gas vesicles, these colonies will not return into the water column. The colonies isolated from the sediment of the North Basin were able to grow in cultured conditions, in the same way as those from the sediment of the South Basin. Therefore, Microcystiscolonies may survive for a long time under stable conditions of low temperature (ca. 8 °C) and darkness, in the sediment of the deep North Basin, accumulating gradually each year.     

Colony formation in the cyanobacterium Microcystis

Morphological evolution from a unicellular to multicellular state provides greater opportunities for organisms to attain larger and more complex living forms. As the most common freshwater cyanobacterial genus, Microcystis is a unicellular microorganism, with high phenotypic plasticity, which forms colonies and blooms in lakes and reservoirs worldwide. We conducted a systematic review of field studies from the 1990s to 2017 where Microcystis was dominant. Microcystis was detected as the dominant genus in waterbodies from temperate to subtropical and tropical zones. Unicellular Microcystis spp. can be induced to form colonies by adjusting biotic and abiotic factors in laboratory. Colony formation by cell division has been induced by zooplankton filtrate, high Pb²⁺ concentration, the presence of another cyanobacterium (Cylindrospermopsis raciborskii), heterotrophic bacteria, and by low temperature and light intensity. Colony formation by cell adhesion can be induced by zooplankton grazing, high Ca²⁺ concentration, and microcystins. We hypothesise that single cells of all Microcystis morphospecies initially form colonies with a similar morphology to those found in the early spring. These colonies gradually change their morphology to that of M. ichthyoblabe, M. wesenbergii and M. aeruginosa with changing environmental conditions. Colony formation provides Microcystis with many ecological advantages, including adaption to varying light, sustained growth under poor nutrient supply, protection from chemical stressors and protection from grazing. These benefits represent passive tactics responding to environmental stress. Microcystis colonies form at the cost of decreased specific growth rates compared with a unicellular habit. Large colony size allows Microcystis to attain rapid floating velocities (maximum recorded for a single colony, ∼ 10.08 m h⁻¹) that enable them to develop and maintain a large biomass near the surface of eutrophic lakes, where they may shade and inhibit the growth of less‐buoyant species in deeper layers. Over time, accompanying species may fail to maintain viable populations, allowing Microcystis to dominate. Microcystis blooms can be controlled by artificial mixing. Microcystis colonies and non‐buoyant phytoplankton will be exposed to identical light conditions if they are evenly distributed over the water column. In that case, green algae and diatoms, which generally have a higher growth rate than Microcystis, will be more successful. Under such mixing conditions, other phytoplankton taxa could recover and the dominance of Microcystis would be reduced. This review advances our understanding of the factors and mechanisms affecting Microcystis colony formation and size in the field and laboratory through synthesis of current knowledge. The main transition pathways of morphological changes in Microcystis provide an example of the phenotypic plasticity of organisms during morphological evolution from a unicellular to multicellular state. We emphasise that the mechanisms and factors influencing competition among various close morphospecies are sometimes paradoxical because these morphospecies are potentially a single species. Further work is required to clarify the colony‐forming process in different Microcystis morphospecies and the seasonal variation in this process. This will allow researchers to grow laboratory cultures that more closely reflect field morphologies and to optimise artificial mixing to manage blooms more effectively.     

Evaluation of Microcystis as food for zooplankton in a eutrophic lake

Four experiments were conducted to evaluate Microcystis as food for zooplankton in Lake Kasumigaura, and the following results were obtained. (1) Moina micrura (Cladocera) showed little growth and no reproduction when the animal was reared with Microcystis cultured in the laboratory. The animal did not grow nor reproduce well when Chlorella was mixed with Microcystis as food. (2) Moina micrura assimilated Microcystis much less than Chlorella when the animal fed on single species of Microcystis or a mixture with Chlorella. (3) Microcystis collected from Lake Kasumigaura could not be utilized by Moina micrura even though the colonies were broken up into edible sizes. However, the alga turned into utilizable food when it was decomposed. (4) No inhibitors of Moina micrura population growth could be found in the non-filtered water of Lake Kasumigaura where Microcystis was blooming heavily. Decomposed Microcystis seemed to be utilized by zooplankton as an important food source in Lake Kasumigaura.     

Grazing on <Emphasis Type="Italic">Microcystis</Emphasis> (Cyanophyceae) by testate amoebae with special reference to cyanobacterial abundance and physiological state

We examined the growth of testate amoebae preying on Microcystis whose physiological states were different in laboratory experiments and a hypertrophic pond. We prepared three experimental systems using water samples dominated by Microcystis aeruginosa: light incubation (control), dark incubation (dark), and light incubation with addition of nitrogen and phosphorus (+NP). In all the systems, the colony density of M. aeruginosa decreased slightly during incubation. Physiological activity of phytoplankton as determined by chlorophyll fluorescence was high and almost constant in the control and +NP systems, whereas it decreased in the dark system. Cell densities of testate amoebae increased in the control and +NP systems, whereas in the dark system they remained low. Thus, growth of the amoebae was low in the systems where physiological activity of Microcystis was low. In a hypertrophic pond, cell density of testate amoebae increased and remained high when M. aeruginosa predominated. Cell density of testate amoebae increased remarkably, simultaneously with the increases in M. aeruginosa colony density and phytoplankton physiological activity. We also found a significant correlation between densities of M. aeruginosa colonies and testate amoebae. We suggested that the physiological activity of Microcystis is one important factor affecting the growth of testate amoebae grazing on Microcystis.     

Environmental factors controlling colony formation in blooms of the cyanobacteria Microcystis spp. in Lake Taihu,China

Nitrogen (N) and phosphorus (P) over-enrichment has accelerated eutrophication and promoted cyanobacterial blooms worldwide. The colonial bloom-forming cyanobacterial genus Microcystis is covered by sheaths which can protect cells from zooplankton grazing, viral or bacterial attack and other potential negative environmental factors. This provides a competitive advantage over other phytoplankton species. However, the mechanism of Microcystis colony formation is not clear. Here we report the influence of N, P and pH on Microcystis growth and colony formation in field simulation experiments in Lake Taihu (China). N addition to lake water maintained Microcystis colony size, promoted growth of total phytoplankton, and increased Microcystis proportion as part of total phytoplankton biomass. Increases in P did not promote growth but led to smaller colonies, and had no significant impact on the proportion of Microcystis in the community. N and P addition together promoted phytoplankton growth much more than only adding N. TN and TP concentrations lower than about TN 7.75–13.95 mg L⁻¹ and TP 0.41–0.74 mg L⁻¹ mainly promoted the growth of large Microcystis colonies, but higher concentrations than this promoted the formation of single cells. There was a strong inverse relationship between pH and colony size in the N&P treatments suggesting CO₂ limitation may have induced colonies to become smaller. It appears that Microcystis colony formation is an adaptation to provide the organisms adverse conditions such as nutrient deficiencies or CO₂ limitation induced by increased pH level associated with rapidly proliferating blooms.     

Simulation of water-bloom formation in the cyanobacterium Microcystis aeruginosa

A mechanism for buoyancy increases in the cyanobacterium Microcystisaeruginosa and the associated formation of surface water-bloomsis presented. The mechanism is based on considering a responsetime in the rate of carbohydrate accumulation. When irradianceincreases, the Microcystis cells may require time to increasetheir rate of carbohydrate accumulation. If irradiance decreasesbefore adjustment, the maximum rate of carbohydrate accumulationis not reached. Colony buoyancy increases during mixing whenthe time scales of the light fluctuations are shorter than theresponse time. To examine the mechanism, a model of Microcystisbuoyancy that incorporates the response time has been coupledwith a hydrodynamics model that simulates mixing. The modelwas applied to a shallow lake to show that a prolonged episodeof intense mixing caused the simulated Microcystis coloniesto become excessively buoyant. Once the mixing subsided, thecolonies accumulated at the surface. Decreases in carbohydratewere reduced in large colonies as their size afforded buoyancyforces that could readily overcome the entraining forces ofthe mixing.     

Modelling spatial distributions of Ceratium hirundnella and Microcystis. in a small productive British lake

The short-term relationships between the spatial distributions of phytoplankton and the environmental conditions of Esthwaite Water, a small eutrophic lake in the English Lake District, UK, were examined using a hydrodynamic model. Spatial distributions of phytoplankton were simulated on two occasions the first, when the population was dominated by dinoflagellates; and the second, when the population was dominated by cyanobacteria.Vertical motility of the dinoflagellate Ceratium hirundinellaand buoyancy of the cyanobacteria Microcystis ssprm.were estimated as functions of irradiance. Water velocity fields were estimated through solving the 3-D Navier–Stokes equations on a finite-volume, unstructured non-orthogonal grid. Simulated circulation patterns of water and phytoplankton were similar to those obtained through field observations. Near-surface drift currents were initiated by wind stress, which then generated return currents along the seasonal thermocline. Aggregations of motile Ceratiumthat existed near the thermocline were pushed upwind by the deep return currents and accumulated at upwelling areas. In contrast, near-surface aggregations of Microcystiswere pushed downwind by the surface currents and accumulated at downwelling areas. Horizontal and vertical phytoplankton distributions resulted from the interaction between the vertical motility of the phytoplankton (dependent upon the light environment) and the velocity vectors at the depths at which the phytoplankton accumulated (dependent upon wind stress and morphometry). Modelling showed that phytoplankton motility and buoyancy greatly affect phytoplankton spatial distributions.     

The effect of temperature on growth characteristics and competitions of <Emphasis Type="Italic">Microcystis aeruginosa</Emphasis> and <Emphasis Type="Italic">Oscillatoria mougeotii</Emphasis> in a shallow,eutrophic lake simulator system

Blue-green algal blooms formed by Microcystis and Oscillatoria often occur in shallow eutrophic lakes, such as Lake Taihu (China) and Lake Kasumigaura (Japan). Growth characteristics and competitions between Microcystis aeruginosa and Oscillatoria mougeotii were investigated using lake simulator systems (microcosms) at various temperatures. Oscillatoria was the superior competitor, which suppressed Microcystis, when temperature was <20°C, whereas the opposite phenomenon occurred at 30°C. Oscillatoria had a long exponential phase (20 day) and a low growth rate of 0.22 day⁻¹ and 0.20 day⁻¹ at 15°C and 20°C, respectively, whereas Microcystis had a shorter exponential phase (2–3 days) at 30°C and a higher growth rate (0.86 day⁻¹). Interactions between the algae were stronger and more complex in the lake simulator system than flask systems. Algal growth in the lake simulator system was susceptible to light attenuation and pH change, and algae biomasses were lower than those in flasks. The outcome of competition between Microcystis and Oscillatoria at different temperatures agrees with field observations of algal communities in Lake Taihu, indicating that temperature is a significant factor affecting competition between Microcystis and Oscillatoria in shallow, eutrophic lakes.     

Nutrient limitation of Microcystis aeruginosa in northern California Klamath River reservoirs

Nutrient limitations were investigated in Copco and Iron Gate Reservoirs, on the Klamath River in California, where blooms of the toxin-producing cyanobacterium Microcystis aeruginosa were first reported in 2005. Nutrient enrichment experiments conducted in situ in June and August, 2007 and 2008, determined responses in phytoplankton biomass, Microcystis abundance and microcystin concentration to additions of phosphorus and different forms of nitrogen (NH₄⁺, NO₃, and urea). Microcystis abundance was determined using quantitative PCR targeting the phycocyanin intergenic spacer cpcBA.Total phytoplankton biomass increased with additions of N both before and during Microcystis blooms, with no primary effects from P, suggesting overall N limitation for phytoplankton growth during the summer season. NH₄⁺ generally produced the greatest response in phytoplankton growth, while Microcystis abundance increased in response to all forms of N. Microcystis doubling time in the in situ experiments was 1.24–1.39 days when N was not limiting growth. The results from this study suggest availability of N during the summer is a key growth-limiting factor for the initiation and maintenance of toxic Microcystis blooms in Copco and Iron Gate Reservoirs in the Klamath River.     

The introduced bivalve <Emphasis Type="Italic">Limnoperna fortunei</Emphasis> boosts <Emphasis Type="Italic">Microcystis</Emphasis> growth in Salto Grande reservoir (Argentina): evidence from mesocosm experiments

In order to assess the effects of the introduced bivalve Limnoperna fortunei on water-column properties of Salto Grande reservoir, experiments were conducted using six 400 L mesocosms: 2 with 100 mussels, 2 with 300 mussels, and 2 controls (without mussels). At 0, 1, 2, 3, 7, 14, 21, 28, and 35 days we measured nutrient and chlorophyll a concentrations, counted and identified the phytoplankton, and estimated the density, size, and number of cells of the colonies of Microcystis spp. Cumulative periphyton growth and total accumulated sediments were assessed in all enclosures at the end of the experiment. Throughout the experiment, in the controls ammonia and phosphates dropped to near zero, whereas in the mesocosms with L. fortunei they increased two- to tenfold. Nitrates decreased in all mesocosms. In the presence of the mussel, chlorophyll a and algal cells dropped until day 3 increasing thereafter, whereas in the controls they increased from day 0. Periphyton growth and sediment accumulation were significantly higher in the mesocosms with mussels that in the controls. Cell density, proportion of colonial cells and colony size of Microcystis spp. increased in all enclosures, but these increases were dramatically (and very significantly) higher in enclosures with 100 and, especially, with 300 mussels, than in the controls. Our results indicate that L. fortunei modifies nutrient concentrations and proportions, and promotes aggregation of solitary Microcystis spp. cells into colonies; both these effects can favor blooms of this often noxious cyanobacteria.     

Size-dependent growth of Microcystis colonies in a shallow,hypertrophic lake: use of the RNA-to-total organic carbon ratio

Microcystis was cultured under standard conditions in BG-11 and M-11 media. Using results of an analysis of RNA and total organic carbon (TOC) content, a significant logarithmic relationship between Microcystis growth rate and the RNA/TOC ratio was described to measure the growth rate. Colonial Microcystis samples collected in a shallow, hypertrophic lake (Lake Taihu, China) during May–November 2012 were divided into six size classes (<75, 75–100, 100–150, 150–300, 300–500, and >500 μm), and the RNA/TOC ratio of each class was analyzed to evaluate differences in growth. The growth rate of colonies in the 150–300-μm size class was highest from May to August, but the growth rate increased along with the increase in colony size from September to November. Our results also indicated that water temperature did not change the relationship between growth rate and colony size, but the growth rate of larger colonies was higher than the growth rate of smaller colonies at conditions of low total nitrogen, low total dissolved phosphorus concentration, and high light intensity. Taken together, these results suggest that large colonial Microcystis possess an advantage that is a consequence of this faster growth at lower nutrient concentrations and high light intensities.     

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.     

Variation in species light acquisition traits under fluctuating light regimes: implications for non‐equilibrium coexistence

Resource distribution heterogeneity offers niche opportunities for species with different functional traits to develop and potentially coexist. Available light (photosynthetically active radiation or PAR) for suspended algae (phytoplankton) may fluctuate greatly over time and space. Species‐specific light acquisition traits capture important aspects of the ecophysiology of phytoplankton and characterize species growth at either limiting or saturating daily PAR supply. Efforts have been made to explain phytoplankton coexistence using species‐specific light acquisition traits under constant light conditions, but not under fluctuating light regimes that should facilitate non‐equilibrium coexistence. In the well‐mixed, hypertrophic Lake TaiHu (China), we incubated the phytoplankton community in bottles placed either at fixed depths or moved vertically through the water column to mimic vertical mixing. Incubations at constant depths received only the diurnal changes in light, while the moving bottles received rapidly fluctuating light. Species‐specific light acquisition traits of dominant cyanobacteria (Anabaena flos‐aquae, Microcystis spp.) and diatom (Aulacoseira granulata, Cyclotella pseudostelligera) species were characterized from their growth–light relationships that could explain relative biomasses along the daily PAR gradient under both constant and fluctuating light. Our study demonstrates the importance of interspecific differences in affinities to limiting and saturating light for the coexistence of phytoplankton species in spatially heterogeneous light conditions. Furthermore, we observed strong intraspecific differences in light acquisition traits between incubation under constant and fluctuating light – leading to the reversal of light utilization strategies of species. This increased the niche space for acclimated species, precluding competitive exclusion. These observations could enhance our understanding of the mechanisms behind the Paradox of the Plankton.     

Individual Microcystis colonies harbour distinct bacterial communities that differ by Microcystis oligotype and with time

Interactions between bacteria and phytoplankton in the phycosphere have impacts at the scale of whole ecosystems, including the development of harmful algal blooms. The cyanobacterium Microcystis causes toxic blooms that threaten freshwater ecosystems and human health globally. Microcystis grows in colonies that harbour dense assemblages of other bacteria, yet the taxonomic composition of these phycosphere communities and the nature of their interactions with Microcystis are not well characterized. To identify the taxa and compositional variance within Microcystis phycosphere communities, we performed 16S rRNA V4 region amplicon sequencing on individual Microcystis colonies collected biweekly via high-throughput droplet encapsulation during a western Lake Erie cyanobacterial bloom. The Microcystis phycosphere communities were distinct from microbial communities in whole water and bulk phytoplankton seston in western Lake Erie but lacked ‘core’ taxa found across all colonies. However, dissimilarity in phycosphere community composition correlated with sampling date and the Microcystis 16S rRNA oligotype. Several taxa in the phycosphere were specific to and conserved with Microcystis of a single oligotype or sampling date. Together, this suggests that physiological differences between Microcystis strains, temporal changes in strain phenotypes, and the composition of seeding communities may impact community composition of the Microcystis phycosphere.     

Coupling of phytoplankton and detritus in a shallow,eutrophic lake (Lake Loosdrecht,The Netherlands)

An oscillating steady state is described of phytoplankton, dominated by Prochlorothrix hollandica and Oscillatoria limnetica, and sestonic detritus in shallow, eutrophic Lake Loosdrecht (The Netherlands). A steady-state model for the coupling of the phytoplankton and detritus is discussed in relation to field and experimental data on phytoplankton growth and decomposition. According to model predictions, the phytoplankton to detritus ratio decreases hyperbolically at increasing phytoplankton growth rate and is independent of a lake's trophic state. The seston in L. Loosdrecht contains more detritus than phytoplankton as will apply to many other lakes. The model provides a basis for estimating the loss rate of the detritus, including decomposition, sedimentation and hydraulic loss. In a shallow lake like L. Loosdrecht detritus will continue to influence the water quality for years.