Hyperscums and the population dynamics of Microcystis aeruginosa

Conceptual models, based on 7 years of data, are constructedto simulate the annual cycle and population dynamics of Microcystisaeruginosa in hypertrophic, warm monomictic Hartbees-poort Dam,South Africa in order to assess the role of hyperscum formation.In Hartbeespoort Dam the large summer planktonic population(mean epilimnion biovolumes of 20–50 mm³ I^–1) andthe low wind speed resulted in the formation of hyperscums (thick,crusted accumulations of floating cyanobacteria at wind-protectedsites) containing up to 50% of the total standing crop for 2–3months in 4 out of 5 years. In years in which hyperscums formedthe post-maximal summer population maintained itself throughoutautumn and into late winter before declining to the annual nadir(>1000 cells ml^–1). When hyperscums did not form, orwere artificially removed, the population fell to similarlylow levels as early as May (autumn) and remained small untilthe spring growth phase began. Microcystis cells decompose inthe upper layers of a hyperscum, but this is not a major lossto the planktonic population. Hyperscums are refuges which helpmaintain large planktonic standing crops during winter whengrowth is not possible but have no effect on the long-term (perennial)survival of Microcystis.     

Structural, physical and chemical characteristics of Microcystis aeruginosa hyperscums from a hypertrophic lake

SUMMARY. 1. This paper examines the structural, physical and chemical characteristics of cyanobacterial hyperscums (floating scums of densely packed eyanobacteria. measuring decimetres in thickness, that are covered by a dry crust of photo-oxidized cells) from hypertrophic Hartbeespoort Dam, South Africa. 2. The hyperscum community was a cyanobacterial-baclcrial association, in which the cyanobacterium Microcystis aeruginosa comprised 98% of the biovolumc, with cell concentrations exceeding 10⁹ ml^?1. 3. The buoyancy mechanism of M. aeruginosa and evaporation at the surface led to increasing compaction of the colonies with declining distance from the surface, and the formation of three distinct, but continuous layers: a 1–2 mm dry surface crust (water content: 14%, chlorophyll a concentration: 3 g l^?1), a 5–10 mm compact layer just below the crust (77% water, 1 g l^?1 chlorophyll a), and a less compact layer (96% water, 200–500 mg l^?1 chlorophyll a) from about 1 cm depth to the bottom, comprising the bulk of the hyperscum. 4. The crust attenuated all the incident light and reduced free gas exchange. Beneath it continuously dark anaerobic, highly reduced conditions prevailed. As the hyperscum aged over 3 months in 1984, at 10 cm depth the pH gradually declined from 6.6 to 5.9, interstitial water ammonia-N concentrations increased from 0.45 to 119 mg l^?1, soluble reactive phosphorus from 2.8 to 83.3 mg l^?1, and dissolved organic carbon reached a maximum of 460 mg l^?1. At any point in time these concentrations declined gradually with increasing depth within the hyperscum, and declined dramatically beneath the hyperscum. Similar patterns were recorded in another hyperscum in 1986. 5. The chemical and temperature depth profiles indicated that free water movement took place around and under the hyperscum, but within it water movement was restricted to diffusion. 6. Gas bubbles composed of 28% methane. 19% CO₂, 53% N₂, and traces of H₂ trapped within the hyperscum, and the presence of volatile fatty acids in the interstitial water were indicative of anaerobic decomposition processes mediated by fermenting and methanogenic bacteria, and N:P ratios below 1.5 in the interstitial water suggested that nitrogen was lost as gas, possibly through denitrification. 7. We hypothesize that the major sites of decay of M. aeruginosa were the crust and the compact layer beneath it, while deeper within the hyperscum this cyanobacterium could survive prolonged periods of dark anaerobic conditions. This hypothesis requires confirmation.     

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.     

Influence of Cyanobacterial Hyperscum on Heterotrophic Activity of Planktonic Bacteria in a Hypertrophic Lake

The response of the planktonic heterotrophic bacterial community to the buildup and breakdown of a semipermanent, crusted, floating cyanobacterial mat, or hyperscum, that covered 1 to 2 ha was studied in a hypertrophic lake (Hartbeespoort Dam, South Africa). The initial response of bacteria in the main basin to the release of dissolved organic carbon (DOC) from the hyperscum 1 km away was an increase in activity per cell from 35 × 10⁻¹² to 153 × 10⁻¹² μg of C cell⁻¹ h⁻¹ for total cell counts, while activity per cell for metabolically active cells increased from 19 × 10⁻¹¹ to 85 × 10⁻¹¹ μg of C cell⁻¹ h⁻¹. No major population growth occurred at this stage. Later, with the continuous supply of DOC from the hyperscum, total bacterial numbers increased from 6.6 × 10⁶ to 20 × 10⁶ cells ml⁻¹, while the activity per cell declined. Metabolically active bacteria followed the same trend. Shorter-term DOC increases caused only increases in bacterial activity per cell. The data from Hartbeespoort Dam demonstrate an interesting and little-documented mechanism by which aquatic bacteria respond to increased DOC concentration and which may be universal for aquatic systems.     

Effects of the Distribution of a Toxic Microcystis Bloom on the Small Scale Patchiness of Zooplankton

Toxic cyanobacterial blooms can strongly affect freshwater food web structures. However, little is known about how the patchy occurrence of blooms within systems affects the spatial distribution of zooplankton communities. We studied this by analysing zooplankton community structures in comparison with the spatially distinct distribution of a toxic Microcystis bloom in a small, shallow, eutrophic lake. While toxic Microcystis was present at all sites, there were large spatial differences in the level of cyanobacterial biomass and in the zooplankton communities; sites with persistently low cyanobacterial biomass displayed a higher biomass of adult Daphnia and higher zooplankton diversity than sites with persistently high cyanobacterial biomass. While wind was the most likely reason for the spatially distinct occurrence of the bloom, our data indicate that it was the differences in cyanobacterial biomass that caused spatial differences in the zooplankton community structures. Overall, our study suggests that even in small systems with extensive blooms ‘refuge sites’ exist that allow large grazers to persist, which can be an important mechanism for a successful re-establishment of the biodiversity in an ecosystem after periods of cyanobacterial blooms.     

CYANOBACTERIAL BUOYANCY REGULATION: THE PARADOXICAL ROLES OF CARBON1

In stratified lakes, dominance of the phytoplankton by cyanobacteria is largely the result of their buoyancy and depth regulation. Bloom-forming cyanobacteria regulate the gas vesicle and storage polymer contents of their cells in response to interactive environmental factors, especially light and nutrients. While research on the roles of nitrogen and phosphorus in cyanobacterial buoyancy regulation has reached a consensus, evaluations of the roles of carbon have remained open to dispute. We investigated the various effects of changes in carbon availability on cyanobacterial buoyancy with continuous cultures of Microcystis aeruginosa Kuetz. emend. Elenkin (1924), a notorious bloom-former. Although CO₂ limitation of photosynthesis can promote buoyancy in the short term by preventing the collapse of turgor-sensitive gas vesicles and/or by limiting polysaccharide accumulation, we found that sustained carbon limitation restricts buoyancy regulation by limiting gas vesicle as well as polysaccharide synthesis. These results provide an explanation for the positive effects of bicarbonate enrichment on cyanobacterial nitrogen uptake and bloom formation in lake experiments and may help to explain the pattern of cyanobacterial dominance in phosphorus-enriched, low-carbon lakes.     

Aspects of the phosphorus cycle in Hartbeespoort Dam (South Africa)

The role of biotic processes in a warm, monomictic, hypertrophic African impoundment (Hartbeespoort Dam) is examined using ³²P radiobioassays. Phosphorus demand is assessed by phosphorus turnover times, alkaline phosphatase activity, cellular phosphorus status and the phosphorus deficiency index. Long turnover times indicative of an enriched system were recorded, ranging from 9 h to 1992 h, with no evidence of phosphorus stress being present. These turnover times support the hypothesis that the phosphorus cycle in Hartbeespoort Dam is dominated by the algal community which is shown to play an important role in phosphorus cycling within the water column. However, hydrological processes remain the driving force in phosphorus seasonality in the lake.     

Role of Predatory Bacteria in the Termination of a Cyanobacterial Bloom

Changes in cyanobacterial abundance and in the occurrence of bacteria of bacteria capable of lysing cyanobacteria were monitored over a period of 6 months (May to October 1998) in eutrophic Brome Lake (Quebec, Canada), in which dense cyanobacterial blooms recur regularly. By screening lake water, we isolated two strains of lytic bacteria, from the family Cytophagaceae. When tested on 12 cyanobacteria and 6 heterotrophic bacteria, strain 1 lysed only Anabaena flos-aquae and strain 2 lysed only Synechococcus cedorum, Synechococcus leopoliensis, Synechococcus elongatus, and Anacystic nidulans: both liquid and agar-grown cultures of these cyanobacteria were lysed. The number of plaque forming units of bacteria increased dramatically during the decline of the bloom. The results are consistent with an important role for these host-specific lytic bacteria in control and elimination of cyanobacterial blooms in this lake.     

Diurnal mixed layers and the long-term dominance of Microcystis aeruginosa

The population dynamics of the cyanobacterium Microcystis aeruginosain a hypertrophic, subtropical lake (Hartbeespoort Dam, SouthAfrica) were followed over 4.5 years. We examined the hypothesisthat M.aeruginosa dominated (>80% by volume) the phytoplanktonpopulation up to 10 months of each year because it maintaineditself within shallow diurnal mixed layers and was thus ensuredaccess to light, the major limiting resource. Wind speeds overHartbeespoort Dam were strong enough to mix the entire epilimnionthrough Langmuir circulations only 12% of the time. At othertimes solar heating led to the formation of diurnal mixed layers(z₁) that were shallower (<2 m) than the euphotic zone (z_cu;mean = 3.5 m, range: 0.45–8.4 m) while the seasonal mixedlayer (z_m) was always deeper than z_cu (range: 7–18 m).By means of its buoyancy mechanism M.aeruginosa maintained thebulk of its population within z₁, while non-buoyant speciessank into dark layers. Adaptation to strong light intensitywas implicated from low cellular chlorophyll a content (0.132µg/106 cells) and high I_k (up to 1230 µE m^–2s^–1). Ensured access to light, the post-maximum summerpopulations persisted throughout autumn and winter, despitesuboptimal temperatures, by sustaining low losses. Increasedsedimentation losses caused a sharp decline of the populationat the end of winter each year, and a short (2–3 months)successional episode followed, but by late spring M.aeruginosawas again dominant. The data from Hartbeespoort Dam point outthe importance of distinguishing between z_m and z₁, and showthe profound effect that the daily pattern of z₁ as opposedto the seasonal pattern of z_m, can have on phytoplankton populationspecies composition.     

Abiotic factors influencing cyanobacterial bloom development in the Gulf of Finland (Baltic Sea)

Blooms of cyanobacteria are a recurrent phenomenon in the Baltic Sea, including the Gulf of Finland. The spatial extension, duration, intensity and species composition of these blooms varies widely between years. Alg@line data collected regularly from ferries as well as weather service and marine monitoring data from 1997 to 2005 are analysed to determine the main abiotic factors influencing the intensity and species composition of cyanobacterial blooms in the Gulf of Finland. It is demonstrated that the development of the Nodularia spumigena Mertens bloom is highly dependent on weather conditions such as photosynthetically active radiation and water temperature. Nutrient conditions, especially the surplus of phosphorus (according to Redfield ratio) related to the pre-bloom upwelling events in the Gulf, affect the intensity of Aphanizomenon sp. (L.) Ralfs blooms. Differences in bloom timing and duration indicate that, if the preconditions (like nutrient ratio/concentration and weather conditions) for bloom formation are favourable, then the Aphanizomenon bloom starts earlier, the overall bloom period is longer and the Nodularia peak might appear in a wider time window. Handling editor: K. Martens     

Diversity and dynamics of microcystin—Producing cyanobacteria in China's third largest lake, Lake Taihu

Microcystins (MC), the most prevalent group of harmful cyanobacterial hepatotoxins, are primarily produced by strains of cyanobacteria in Microcystis, Anabaena and Planktothrix. Lake Taihu, which is the third largest freshwater lake in China, is a hypertrophic shallow lake in eastern China that has experienced lake-wide cyanobacterial blooms annually during the last few decades. In this study, PCR-DGGE was used to evaluate the diversity of potential MC-producing cyanobacteria and real-time PCR was used to analyze the dynamics of this population based on the presence of the mcy gene in samples collected during a year long study. The results revealed that all MC-producing genotypes detected belonged to the genus Microcystis. In addition, the MC-producing genotype communities were more diverse during the bloom season than the non-bloom season, and the diversity in the late bloom period was lower than the diversity in the early bloom period. Furthermore, the abundance of MC-producing genotypes increased dramatically during the bloom development period, reaching its peak in late summer (September). The results also suggested that the highest mcy gene concentration lagged behind the highest MC concentration, and the potential MC-producing cyanobacterial community shift lagged behind the development of blooms.     

Increased risk of cyanobacterial blooms in northern high‐latitude lakes through climate warming and phosphorus enrichment

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Biological and chemical factors driving the temporal distribution of cyanobacteria and heterotrophic bacteria in a eutrophic lake (West Lake,China)

Physico-chemical parameters, hydrological conditions, and microbial interactions can affect the growth and persistence of cyanobacteria, but the interacting effects among these bloom-forming factors are still poorly known. This hampers our capacity to predict the occurrence of cyanobacterial bloom accurately. Here, we studied the relationship between temperature, N and P cycles, and the microbial community abundance and diversity at 0.5 m under the surface of West Lake (China) from January 21 to November 20, 2015, in order to better understand the key factors regulating temporal changes in the cyanobacterial community. Using high throughput sequencing of the 16S rRNA gene V3-V4 region, we studied the diversity and abundance of bacteria. In parallel, we measured physico-chemical parameters and followed the abundance of key genes involved in N fixation, denitrification, and nutrient uptake. Multivariate analyses suggest that P concentration and water temperature are the key factors controlling the outbreak of summer cyanobacterial bloom. RT-qPCR analyses of the bacterial community and measurements of the copy number of denitrification-related gene (nirK, nosZ, nirS) show that denitrification potential and denitrifying bacteria relative abundance (Pseudomonas and Bacillus) increased in concert with diazotrophic cyanobacterial genera (Anabaena, Nostoc, Aphanizomenon flos-aquae) and the common bloom-forming non-diazotrophic cyanobacterium genus Microcystis. The present study brings new insights on the complex interplay between physico-chemical parameters, heterotrophic bacterial community composition, nitrogen cycle, and cyanobacteria dominance in a eutrophic lake.

     

Cyanobacterial dominance in lakes

Cyanobacterial dominance in lakes has received much attention in the past because of frequent bloom formation in lakes of higher trophic levels. In this paper, underlying mechanisms of cyanobacterial dominance are analyzed and discussed using both original and literature data from various shallow mixed and deep stratifying lakes from temperate and (sub)tropical regions. Examples include all four ecotypes of cyanobacteria sensu Mur et al. (1993), because their behavior in the water column is entirely different. Colony forming species (Microcystis) are exemplified from the large shallow Tai Hu, China. Data from a shallow urban lake, Alte Donau in Austria are used to characterize well mixed species (Cylindrospermopsis), while stratifying species (Planktothrix) are analyzed from the deep alpine lake Mondsee. Nitrogen fixing species (Aphanizomenon) are typified from a shallow river-run lake in Germany. Factors causing the dominance of one or the other group are often difficult to reveal because several interacting factors are usually involved which are not necessarily the same in different environments. Strategies for restoration, therefore, depend on both the cyanobacterial species involved and the specific causing situation. Some uncertainty about the success of correctives, however, will remain due to the stochastic nature of the events and pathways leading to cyanobacterial blooms. Truly integrated research programs are required to generate predictive models capable of quantifying key variables at appropriate spatial and temporal scales.     

Longitudinal Differences of Phytoplankton Community during a Period of Small Water Level Fluctuations in a Subtropical Reservoir Bay (Xiangxi Bay,Three Gorges Reservoir,China)

The Three Gorges Reservoir (TGR) had a great inflow discharge with small water level fluctuations in the flood season of the year 2008, when a large‐scale cyanobacterial bloom broke out in Xiangxi Bay (a tributary bay behaving like a lake) for the first time after the construction of the TGR impoundment. To compare spatiotemporal longitudinal differences of phytoplankton community structure during this period, weekly surveys were performed in Xiangxi Bay. The cyanobacterial bloom lasted from June 6 to July 18, 2008, with Microcystis aeruginosa as the dominant species. During this bloom the species diversity, evenness and community change rate was relatively low. The probable causes for the interruption of the bloom were precipitation and water temperature. In the non‐cyanobacterial bloom period (July 25 to September 26, 2008) many other species dominated the community including Stephanodiscus hantzschii and Cryptomonas ovata, with higher values of species diversity, evenness and community change rate. As for the longitudinal differences, the community structure in the riverine zone was different from that in other zones, indicating the important effect of inflow from the upstream of Xiangxi Bay. The management actions in Xiangxi Bay should prevent blooms in the mainstream, lacustrine and transitional zone. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Effects of Calcium Levels on Colonial Aggregation and Buoyancy of <Emphasis Type="Italic">Microcystis aeruginosa</Emphasis>

Colonial aggregation of Microcystis plays a key role in bloom formation. Limited studies have been reported about effects of environmental factors on the aggregation of Microcystis. Calcium is an important chemical element in water system. In this study, we investigated the effects of a low- (0.015 g l⁻¹) and a high-concentration of calcium (0.100 g l⁻¹) on the aggregation and buoyancy of a colonial strain M. aeruginosa XW01. Results show that compared to the low concentration of calcium, the high-calcium condition results in bigger colonial size, higher level of buoyancy and increased production of extracellular polysaccharides (EPS) of M. aeruginosa XW01. Increased production of EPS induced by the high-calcium concentration should contribute to the colonial aggregation and buoyancy of M. aeruginosa XW01. These results suggest that an increase in calcium concentration may be beneficial for Microcystis blooms occurring in a soft water lake.     

To bloom or not to bloom: contrasting responses of cyanobacteria to recent heat waves explained by critical thresholds of abiotic drivers

Past heat waves are considered harbingers of future climate change. In this study, we have evaluated the effects of two recent Central European summer heat waves (2003 and 2006) on cyanobacterial blooms in a eutrophic, shallow lake. While a bloom of cyanobacteria developed in 2006, consistent with our expectations, cyanobacterial biomass surprisingly remained at a record-low during the entire summer of 2003. Critical thresholds of abiotic drivers extracted from the long-term (1993–2007) data set of the studied lake using classification tree analysis (CTA) proved suitable to explain these observations. We found that cyanobacterial blooms were especially favoured in 2006 because thermal stratification was critically intense (Schmidt stability >44 g cm cm⁻²) and long-lasting (>3 weeks). Our results also suggest that some cyanobacterial species (Anabaena sp.) benefitted directly from the stable water column, whereas other species (Planktothrix sp.) took advantage of stratification-induced internal nutrient loading. In 2003, conditions were less favourable for cyanobacteria due to a spell of lower temperatures and stronger winds in mid-summer; as a result, the identified thresholds of thermal stratification were hardly ever reached. Overall, our study shows that extracting critical thresholds of environmental drivers from long-term records is a promising avenue for predicting ecosystem responses to future climate warming. Specifically, our results emphasize that not average temperature increase but changes in short-term meteorological variability will determine whether cyanobacteria will bloom more often in a warmer world.     

Toxin-Producing Anabaena flos-aquae Induces Settling of Chlamydomonas reinhardtii,a Competing Motile Alga

Toxin production is an adaptation that allows cyanobacteria in resource-limiting environments to ameliorate the effects of herbivory and competition with other phototrophs. We demonstrate that the cyanobacterial toxins anatoxin-a and microcystin-LR paralyze the motile green alga Chlamydomonas reinhardtii. In addition, both purified toxins and cyanobacterial extracellular products containing these toxins cause the alga to settle faster than in nontoxic media. In microcosm experiments, the presence of either the cyanobacterium or its extracellular products induce settling in the alga, similar to the response observed with the addition of both anatoxin-a and microcystin-LR. The cyanobacterial production of paralyzing toxins represents a novel mechanism for phytoplankton settling. This prokaryotic/eukaryotic chemical interaction may create a competitor-free zone for cyanobacteria in lake environments, predicating optimal conditions for a toxic cyanobacterial bloom.     

A new biomonitoring protocol to determine the ecological health of impoundments using artificial substrates

The colonisation potential of macroinvertebrate families was measured using artificial substrates. These substrates were tested on three highveld impoundments. Bronkhorstspruit Dam was an amictic, mesotrophic impoundment, while Hartbeespoort and Roodeplaat Dams were both monomictic, eutrophic impoundments. Bronkhorstspruit Dam had the best water quality and the smallest algal population, comprising mainly diatom species. Hartbeespoort Dam had the most saline water (TDS 96–400 mg/l; NH₄ <0.004–0.218mg/l; NO₂+NO₃ 0.109–2.776mg/l; PO₄ <0.04–0.06mg/l), while Roodeplaat Dam had the highest nutrient concentrations (TDS 200–373mg/l; NH₄ <0.04–0.933mg/l; NO₂+NO₃ <0.04–2.310mg/l; PO₄ 0.041–0.472 mg/l). Both Hartbeespoort and Roodeplaat Dams had large algal populations that resulted in very alkaline water during summer. Microsystis dominated the phytoplankton population in these two impoundments during summer and autumn, while green algae and diatoms were dominant during winter and spring. Ecological health was determined using two different biotic indices. Two modifications of the Belgian Biotic Index and a modification of SASS4 (SASSD) were tested. Three health classes were suggested for SASSD scores and ASPT values. It is concluded that artificial substrates with SASSD (and ASPT) as a biotic index can be used to determine the biological health of impoundments, but that further refinement of the index is needed.     

The Stream Inlet to a Shallow Bay of a Drinking Water Reservoir,a ‘Hot‐Spot’ for Microcystis Blooms Initiation

Stream inlets into shallow bays of reservoirs and lakes may be ‘hot‐spots’ for toxic cyanobacterial bloom initiation. These ‘hot‐spots’ may be connected with the permanent inflow of high nutrient concentrations from the catchment, optimal physical conditions (wind protected areas) that occur in shallow areas and/or ineffective top‐down control. Four sampling sites along a transect from stream to reservoir in a shallow bay of Sulejow Reservoir (Poland) were studied to test the above hypothesis, comprising a transition zone between lotic and pelagic habitats. Investigations showed that stream inlet into shallow bay acted as incubator for Microcystis blooms. The nutrient level, especially phosphorus, was identified as the major cause of cyanobacterial bloom growth. The increase of Microcystis biomass strongly correlated with increasing microcystin concentrations, however, a relationship with microcystin content was not observed. Toxicity of bloom demonstrated seasonal variability, reaching its maximum at the initial phase of bloom. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)