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.     

Buoyancy regulation in light-limited continuous cultures of Microcystis aeruginosa

Microcystis aeruginosa was grown in light-limited continuouscultures at different growth rates on light-dark cycles at variousphotopenods. Due to the strength of the gas vesicles the organismwas not able to collapse its gas vesicles by turgor pressure.Below the maximal growth rate, the organism was buoyant dueto its high gas vesicle content. The results suggested thatthe rate of gas vesicle synthesis was not regulated. Upon atransition to high irradiance it took several hours before thecells lost their buoyancy due to polyglucan accumulation. Theresults are interpreted in an ecological context and it is suggestedthat Microcystis is an epilimnetic species due to its buoyancyregulation.     

Diurnal buoyancy changes of Microcystis in an artificially mixed storage reservoir

In a storage reservoir, which is artificially mixed in order to reduce algal and especially cyanobacterial growth, the cyanobacterium Microcystis is still present. The aim of the research was to investigate why Microcystis was able to grow in the artificially mixed reservoir. From the results it could be concluded that the large shallow area in the reservoir allows this growth. The loss of buoyancy during the day was much higher in this shallow part than in the deep part. Assuming that the loss of buoyancy was the result of a higher carbohydrate content, a higher growth rate in the shallow part may be expected. A higher received light dose by the phytoplankton in the shallow mixed part of the reservoir than in the deep mixed part explains the difference in buoyancy loss. A significant correlation between the received light dose (calculated for homogeneously mixed phytoplankton) and the buoyancy loss was found. Apparently, the Microcystis colonies were entrained in the turbulent flow in both the shallow and the deep part of the reservoir. With a little higher stability on one sampling day, due to the late start of the artificial mixing, the loss of buoyancy at the deep site was higher than on the other days and almost comparable to the loss at the shallow site. Although the vertical biomass distribution and the temperature profiles showed homogeneous mixing, the colonies in the upper layers apparently received a higher light dose than those deeper in the water column. Determination of the buoyancy state of cyanobacteria appeared to be a valuable method to investigate the light history and hence their entrainment in the turbulent flow in the water column.     

Role of light, carbon dioxide and nitrogen in regulation of buoyancy, growth and bloom formation of Anabaena flos-aquae

The availability of light, CO₂ and NH₄-N interacted to controlbuoyancy and growth of the gas vacuolate blue-green alga, Anabaenaflos-aquae. At high light intensities algal growth rates werehigh; however, the alga was non-buoyant regardless of the availabilityof CO₂ or NH₄-N. The mechanism for buoyancy loss involved increasedcell turgor pressures at higher light intensities which resultedin collapse of gas vacuoles. At lower light intensities algalgrowth rates and cell turgor pressures were reduced and buoyancywas controlled by the availability of CO₂ and inorganic nitrogen.Carbon dioxide limitation increased buoyancy, while reducedinorganic nitrogen availability reduced buoyancy. Mechanismsfor buoyancy regulation at low light intensities involved changesin cellular C/N ratios which appeared to affect the rate ofsynthesis and accumulation of protein-rich gas vacuoles. Algalspecific growth rates were combined with buoyancy data to forma single index (µ_bloom) to the rate of surface bloom formationof A.flos-aquae as a function of the availability of light,CO₂ and NH₄-N. The bloom formation index was enhanced with decreasedavailability of light and CO₂, and increased availability ofNH₄-N.     

Autunmal sedimentation of Microcystis spp. as result of an increase in carbohydrate ballast at reduced temperature

Autumnal sedimentation of the Microcystis population was studiedin Lake Nieuwe Meer (Amsterdam, The Netherlands). In summer,Microcystis formed a high percentage of the total phytoplanktonin the water column, but a low percentage in sedimentation traps.The reverse was found during September and October, with a highpercentage in the sedimentation traps, but a low percentagein the water column. The decrease in the numbers of Microcystiscolonies coincided with a decrease in waler temperature. Inexperiment with a strain of Mirocystis, isolated from Lake NieuweMeer, the percentage of total colonies that were sinking increasedin a few days to 100% after a shift from 20C to 15.3, 13.0or 10.5C. The gas vesicle volume in the cells remained constantduring the incubations. Sinking of the colonies resulted froman increased glycogen content. Calculation of carbon (C) flowsduring the first 2 days of the incubation at reduced temperatureshowed that the glycogen accumulation was the result of a muchlower rate of protein synthesis during the light period at thelower temperatures. Although the photosynthetic rate itselfdecreased at reduced temperature, it resulted in more fixedCO₂ being stored as glucose. Because the respiratory rate alsodecreased (with an almost similar decrease to that of photosynthesis),glycogen accumulated at lower temperatures. It was calculatedthat after an incubation period of-I week at reduced temperature,the rate of photosynthesis had decreased by 10.1% of the valueat 20C per 1C, while the rate of respiration had decreasedonly 1.8%. It is proposed that there is a feedback mechanismin which an increasing concentration of glycogen inhibits photosynthesisand stimulates respiration.     

Uptake of 13C and 15N (ammonium, nitrate and urea) by Microcystis in Lake Kasumigaura

The uptake rate of carbon and nitrogen (ammonium, nitrate andurea) by the Microcystis predominating among phytoplankton wasinvestigated in the summer of 1984 in Takahamaira Bay of LakeKasumigaura. The V_max values of Microcystis for nitrate (0.025–0.046h^–1) and ammonium (0.15–0.17 h^–1) were considerablyhigher than other natural phytoplankton. The ammonium, nitrateand urea uptake by Microcystis was light dependent and was notinhibited with nigh light intensity. The K₁ values were farlower than the I_k values. The carbon uptake was not influencedby nitrogen enrichment. Microcystis accelerated the uptake rateby changing V_max/K _s value when nitrogen versus carbon contentin cells declined. Nitrate was scarcely existent in TakahamairiBay during the summer, when Microcystis usually used ammoniumas the nitrogen source. However, the standing stock of ammoniumin the water was far lower than the daily ammonium uptake rates.Therefore, the ammonium in this water had to be supplied becauseof its rapid turn-over time (–0.7–2.6 h).     

Hyperscums of the cyanobacterium Microcystis aeruginosa in a hypertrophic lake (Hartbeespoort Dam, South Africa)

It is proposed that surface scums of densely packed planktoniccyanobacteria (blue-green algae) which exist for weeks to months,measure several decimeters in thickness and are covered by acrust of photo-oxidized cells, be called hyperscums. Hyperscumsof Microcystis aeruginosa formed during prolonged periods ofcalm weather in wind-protected sites in a hypertrophic lakesubject to low wind speeds (Hart beespoort Dam, South Africa).A hyperscum that extended over 1–2 hectares and persistedfor 103 days during winter 1983 was studied. Chlorophyll a concentrationsranged from 100 to 300 mg l^–2 Microcystis cell concentrationsreached 1.76x10⁹ cells ml^–1 or 116 cm³l^–1. The hyperscumenvironment was anoxic, aphotic, with a fluctuating temperatureregime and low pH values. The densely packed Microcystis cellssurvived these conditions for more than 2 months. This was shownby comparing the potential photosynthetic capacity of Microcystisfrom the hyperscum with that of Microcystis from the main basinof the lake. However, after 3 months the hyperscum algae losttheir photosynthetic capacity and decomposition processes prevailed.The hyperscum gradually shrank in size until a storm causedits complete disintegration.     

Effects of macronutrients upon buoyancy regulation by metalimnetic Oscillatoria agardhii in Deming Lake, Minnesota

Gas-vacuolate filaments of Oscillatoria agardhii form a metalimneticlayer in Oeming Lake, Minnesota. The environmental factors whichaffect buoyancy and the physiological processes which mediatechanges in buoyancy were determined. Buoyant filaments losttheir buoyancy in a few hours when incubated at light intensitiesabove those found in situ ({small tilde}15 µnol photonsm^–2 s^–1, or 1% of the surface value). The rate ofbuoyancy loss was accelerated by the addition of 10 µMphosphate at irradiances >200mol photons m^–2 s^–1.The effect of nutrient additions on buoyancy was also investigatedover a longer time period by incubating metalimnetic samplesin situ. The samples were deployed for 6 days at a depth wherethe irradiance was 8% of the surface value. As found in short-termexperiments, the addition of phosphate resulted in the largestdecrease in buoyancy. However, the addition of ammonia in additionto phosphate attenuated the buoyancy loss on day 2, and on day6 the filaments in these treatments were almost completely buoyant.The physiological status of the filaments in these treatmentswas assayed by analysis of elemental ratios of C, N and P, andby measurement of cellular chlorophyll, polysaccharide and protein.In addition, the cellular content of gas vesicles was determined.The construction of ballast balance sheets from these data indicatedthat changes in buoyancy were primarily due to differences inthe amount of polysaccharide ballast in the cells. However,in another set of in situ experiments, the increase in measuredballast molecules did not explain the observed loss of buoyancy.We hypothesized that another, undetected ballast-providing moleculehad accumulated in the cells.     

Resource quality effects on Daphnia longispina offspring fitness

The present study addressed the question: is the fitness ofDaphnia longispina neonates related to the quality (species)of the resource their mothers consumed? A 2 2 factorial designwith two maternal and neonate resource types was employed, withRhodomonas and Microcystis used as high- and low-quality resources,respectively. Neonate fitness was assessed using demographicparameters obtained from a life-table study. The resource typeneonates consumed always had a stronger effect on the neonatesthan did the maternal type. Neonate consumption of Rhodomonas.relative to Microcystis. increased fitness, as estimated bythe instantaneous population growth rate, by 25–28%. whilematernal consumption of Rhodomonas increased fitness by 6-8%.Neonate Rhodomonas consumption also reduced the age at reachingthe primiparous instar; increased brood size at the primiparousinstar and body length immediately after the primiparous instar;increases average dutch size, number of clutches produced, totalegg production. and mean body length up to and on age 30 days.Regardless of resource type, maternal Rhodomonas consumptionresulted in neonates which reached maturity at an earlier ageand had a larger individual size immediately after reachingmaturity. The mean clutch size, total egg production, and lengthup to and at age 30 days were all significantly affected bymaternal diet when neonates consumed Microcystis; however, thesevariables were not influenced by maternal type when neonatesconsumed Rhodomonas. This is the first study to document maternallymediated effects of resource quality on the fitness of crustaceanzooplankton neonates.     

Photosynthesis and primary production of Microcystis aeruginosa Kutz. in Lake Kasumigaura

Seasonal changes in the photosynthesis and primary productionof Microcystis aeruginosa Kütz. were investigated in LakeKasumigaura during 1981–1982. Microcystis always showeda light-saturated photosynthesis-light curve. Both P_max andthe initial slope of the photosynthesis-light curve of Microcystisin early summer were very high, so it was concluded that Microcystisutilized both low and high light intensities efficiently. TheP_max of Microcystis was found to be a function of the watertemperature except in August and September. The linear regressionon the temperature-P_max relationship discontinued at 11°C,where the P_max value dropped; Microcystis did not photosynthesizebelow 4°C. The initial slope of the curve was also descendingbelow 11°C. It is suggested that Microcystis changes itsphysiological properties below 11°C. The highest value ofgross production calculated for M. aeruginosa was 5.4 gC m^–2d^–1 in July; the annual gross production was estimatedto be 300 gC m^–2year^–1 (i.e., 40% of the total primaryproduction in this lake).     

The inverse correlation between growth rate and cell carbohydrate content of <Emphasis Type="Italic">Microcystis aeruginosa</Emphasis>

The cell carbohydrate content of cyanobacteria can alter buoyancy, and the ability to regulate the buoyancy is one of the most important mechanisms of cyanobacterial blooms. The net accumulation of carbohydrate in cell is affected by photosynthesis, respiration, synthesis of proteins, and other metabolisms, which are connected to the growth. The aim of this work is to seek the relationship between growth rate and intracellular carbohydrate content. The cell carbohydrate content in Microcystis aeruginosa cultures with different growth characteristics was investigated, and the relationship between growth rate and accumulated carbohydrate in cyanobacterial cells was analyzed. The result showed that the specific growth rate was inversely proportional to cell carbohydrate content. The growth rate was relatively high when the cell carbohydrate content was low. It can be indicated that high growth occurs when cells are buoyant, which favors blooms.     

Sedimentation of phytoplankton populations dominated by Microcystis in a shallow lake

The sedimentary flux of phytoplankton was measured using sedimenttraps in a shallow hypertrophic lake (Lake Kasumigaura), whereMicrocystis bloomed, from June to November 1983 The sedimenttraps were set at 0.5, 1.5 and 3.0 m depth in Takahamairi Bay(3.5 m depth). Microcystis spp. (including M.aerugmosa and M.viridis)in the traps were rare until early August, but increased thereafter.Sinking rates of Microcystis were 0.0045, 0.020 and 0.24 m day^–1in June–August, September and October respectively, whichwere far lower than those of Melosira (0.2–1.7 m day^–1)and Syncdra (0.2–1.0 m day^–1). The total sedimentaryfluxes of POC and that of algal carbon during the study periodwere 283.2 and 96.7 gC m^–2 which were 59.5% and 20.3%of the gross primary production (475.8 gC m^–2) respectively.The sedimentary flux of living algae measured by algal countswas large in June but small in August and September. On theother hand, the flux of detritus obtained by subtracting totalalgal carbon from POC was small in June and July but large inAugust and September. Therefore diatoms, which appeared mostlyin June, tended to sink as live algae, while Microcystis sankas detritus after being decomposed or consumed in the waterIt was concluded from the results of carbon budget calculationsand the respiration rate of the 1- to 20-µm fraction thatthe activity of decomposers or consumers increased greatly inthe short period at the end of the bloom of Microcystis.     

Differential Inhibition of Shootvs.Root Growth at Low Temperature and its Relationship with Carbohydrate Accumulation in Different Wheat Cultivars

Shoot and root growth rate, carbohydrate accumulation (includingfructan), reducing sugar content and dry matter percentage weremeasured in six wheat cultivars, ranging from winter to springtypes, grown at either 5 or 25 °C. At 5 °C (comparedwith 25 °C), the relative growth rate (RGR) of shoots wassimilarly reduced in all cultivars, but the RGR of shoots wasmore affected in winter wheats. This difference resulted insmaller root:shoot ratios than in spring wheats, which alsodeveloped more first-order lateral roots. A direct relationshipbetween carbohydrate accumulation at low temperatures and reductionin root growth was established. These results suggest that differentialshootvs.root growth inhibition at low temperature may play akey role in carbohydrate accumulation at chilling temperatures.This differential response might lead to improvements in survivalat temperatures below 0 °C, regrowth during spring, andwater and nutrient absorption at low temperatures.Copyright1997 Annals of Botany Company Wheat; Triticum aestivum; low temperatures; root growth; root: shoot ratio; sugar accumulation     

Root Respiration and Carbohydrate Status of Two Wheat Genotypes in Response to Hypoxia

To investigate root respiration and carbohydrate status in relationto waterlogging or hypoxia tolerance, root respiration rateand concentrations of soluble sugars in leaves and roots weredetermined for two wheat (Triticum aestivum L.) genotypes differingin waterlogging-tolerance under hypoxia (5% O₂) and subsequentresumption of full aeration. Root and shoot growth were reducedby hypoxia to a larger extent for waterlogging-sensitive Coker9835. Root respiration or oxygen consumption rate declined withhypoxia, but recovered after 7 d of resumption of aeration.Respiration rate was greater for sensitive Coker 9835 than fortolerant Jackson within 8 d after hypoxia. The concentrationsof sucrose, glucose and fructose decreased in leaves for bothgenotypes under hypoxia. The concentration of these sugars inroots, however, increased under hypoxia, to a greater degreefor Jackson. An increase in the ratio of root sugar concentrationto shoot sugar concentration was found for Jackson under hypoxicconditions, suggesting that a large amount of carbohydrate waspartitioned to roots under hypoxia. The results indicated thatroot carbohydrate supply was not a limiting factor for rootgrowth and respiration under hypoxia. Plant tolerance to waterloggingof hypoxia appeared to be associated with low root respirationor oxygen consumption rate and high sugar accumulation underhypoxic conditions.Copyright 1995, 1999 Academic Press Oxygen consumption rate, sugar accumulation, Triticum aestivum L., waterlogging tolerance     

Cladoceran filtration rate-body length relations: model improvements developed for a Microcystis-dominated hypertrophic reservoir

Filtering rates were measured for zooplankton species in Situon single-celled Chlorella and on four Microcystis colony sizefractions (5–20, 20–40, 40–60 and 60–100µm) in a hypertrophic reservoir. Natural-log-transformedfiltration rates of five cladoceran species, one copepod andone rotifer were included in an all-food-particle, all-speciesmultiple regression model which explained 43% of the variancein filtration rate as a function of animal body length. An additional14% and 7.6% of the variance was attributable to food type andzooplankton species respectively, with temperature accountingfor <4% of the variance. Restricting the filtration ratemodel to cladocerans alone explained 51% of the variance asa function of animal length, 16% as a function of food type,7.5% as a function of species and only 0.2% as a function oftemperature. In linear filtration rate models for each foodtype, cladoceran body length explained 70% of the variance whenfeeding on Chlorella and between 57 and 67% of the varianceon the four Microcystis colony fractions. Models describingcladoceran filtration rates on Chlorella and the 5–20µm Microcystis colony fraction were significantly differentfrom the three models on larger colonies due to cladoceran responsesto increasing food particle size. Accordingly, a combined modelfor Microcystis colonies >20 µm was developed. Inclusionof food quality factors such as cyanophyte colony size seemsjustified in models aimed at estimating clearance rates, resourceutilization and phytoplankton grazing losses in plankton orecosystem studies when applied to eutrophic or hypertrophiclakes where large cyanophyte particles are abundant.     

The kinetics of algal photoadaptation in the context of vertical mixing

The responses of phytoplankton to turbulent motions in the surfacemixed layer can be measured to estimate the rate of verticalmixing. If the time scale for the response (photoadaptation)is shorter than that for vertical mixing, phytoplankton willexhibit a vertical gradient associated with adaptation to ambientlight, whereas if mixing occurs with a time scale shorter thanthat of photoadaptation, the surface mixed layer will be uniformwith respect to the photoadaptive parameter. To examine thephysiological bases for a model of vertical mixing and photoadaptation,we grew the marine diatom Thalassiosira pseudonana (clone 3H)at three photon flux densities and subjected the cultures toreciprocal light shifts, measuring physiological and chemicalchanges over the following 10 h. Several parameters, easilymeasured in nature and attributable primarily to phytoplankton,responded to fluctuating light on different time scales. Aftercultures were exposed to relatively bright light, both the initialslope of the photosynthesis-irradiance curve and in vivo fluorescencewere depressed on a time scale of less than an hour. Photosyntheticcapacity was also reduced transiently, but recovered over manyhours to a high level characteristic of an adapted state. First-orderkinetics (the current model of choice for describing photoadaptation)reasonably described the rapid responses of phytoplankton tobright light, but other parameters (i.e. cellular chemical compositionand photosynthetic capacity) changed as a result of unbalancedgrowth and required much longer to adapt from low to high lightas compared to from high to low light. A logistic model of thisadaptation is presented. The model suggests that hysteresisof adaptation during vertical mixing may have important consequences.The vertical distributions of photoadaptive properties in mixedlayers not only reveal the rate of vertical mixing, but showhow phytoplankton integrate environmental fluctuations.     

Variation in life history responses of Daphnia to toxic Microcystis aeruginosa

Three clones of Daphnia pulex and two clones of Daphnia longispinawere exposed to toxic Microcystis aeruginosa for 21 days ina lifetable experiment. The growth and reproduction of individualdaphnids were followed daily to study the long-term effectsof toxic Microcystis. Exposure to Microcystis increased mortality,decreased growth, delayed maturation and decreased offspringproduction, indicating nutritional deficiency and toxic effects.We found variation in life history responses between speciesand among clones. Our results suggest that toxic cyanobacteriamay act as a modifying agent in zooplankton communities at boththe species and clonal level.     

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.