The formation and degradation of cyanobacterium Aphanizomenon flos-aquae blooms: the importance of pH, water temperature, and day length

Investigation of annual changes in phytoplankton community structure in a small artificial eutrophic pond was carried out from May 2002 to April 2003. A heavy bloom of Aphanizomenon flos-aquae var. klebahnii Elenk. (Cyanobacteria) persisted in most of the water column from June to the end of October. In November, the A. flos-aquae bloom suddenly crashed and green algae were predominant until the end of spring. Weekly monitoring suggested strong involvement of the changes in abiotic factors in the cyanobacterial bloom degradation. To clarify the effects of pH, water temperature, and day length on the growth of A. flos-aquae, laboratory batch experiments were conducted. The results showed that A. flos-aquae could not grow below pH 7.1 and 11°C, and the growth tended to be suppressed under a 10L:14D photoperiod. pH, water temperature, and day length are vital factors in the growth of A. flos-aquae and, additionally, grazing by cyclopoid copepods also seemed important in bloom collapse.     

Disturbance events affecting phytoplankton biomass,composition and species diversity in a shallow,eutrophic, temperate lake

The seasonal changes in phytoplankton biomass and species diversity in a shallow, eutrophic Danish lake are described and related to different disturbance events acting on the phytoplankton community.Both the spring diatom maximum and the summer bloom of the filamentous blue-green alga, Aphanizomenon flos-aquae (L.) Ralfs, coincided with low values of phytoplankton species diversity and equitability. Diatom collapse was mainly due to internal modifications as nutrient depletion (Si, P) caused by rapid growth of phytoplankton, and increased grazing activity from zooplankton. A large population of Daphnia longispina O.F. Müller in June effectively removed smaller algal competitors, thus favouring the development of a huge summer bloom (140 mm³ l^–1) of Aphanizomenon flos-aquae. Heavy rainfall and storms in late July increased the loss of Apahnizomenon by out-flow and disturbed the stratification of the lake. These events caused a marked decline in phytoplankton biomass but had no effect on species diversity. A second storm period in late August circulated the lake completely and was followed by a rapid increase in phytoplankton diversity, and a change in the phytoplankton community structure from dominance of large, slow-growing K-selected species (Aphanizomenon) to small, fast-growing r-selected species (cryptomonads).     

浅水湖泊浮水植物芡实收割对浮游植物群落的影响

浅水湖泊生态系统正遭受广泛而强烈的人为干扰,但是对收割水生植物干扰的研究甚少。于2019年8月对芡实过度生长的陈瑶湖进行通道式分区收割工程,分析了收割芡实（Euryale ferox）前后不同处理组浮游植物群落的变化。研究期间共鉴定出浮游植物6门47属72种,其中收割前63种,收割后71种。收割后浮游植物的细胞密度和生物量均高于收割前,分别增加了39.78%和5.09%。收割芡实导致陈瑶湖浮游植物群落为由蓝藻-绿藻-硅藻-隐藻群落转变为蓝藻-绿藻-硅藻群落。其中蓝藻细胞密度和生物量显著高于收割前（P<0.05）,归因于有害蓝藻（铜绿微囊藻Microcystis aeruginosa、水华束丝藻Aphanizomenon flos-aquae、小颤藻Oscillatoria tenuis、卷曲鱼腥藻Dolicospermum circinale、小席藻Phormidium tenu）的增加。收割还导致了硅藻群落由附生型向浮游型硅藻的转变,表现为尖针杆藻（Ulnaria acus）减少,而颗粒直链藻极狭变种（Aulacoseira granulata var.angustissima）、梅尼小环藻（Cyclotella meneghiniana）增加。在芡实收割过程中,未收割组和河道的浮游植物群落结构在时空分布上无显著性差异（P>0.05）,但收割组在收割后的不同阶段内差异较为明显,其细胞密度和生物量随着收割实验的结束逐渐降低。浮游植物与环境因子的相关性分析表明,水生植被覆盖度、总磷、总氮、溶解氧和叶绿素a浓度是影响浮游植物细胞密度和生物量变化的主要环境因子。综合陈瑶湖水质状态,本研究认为收割芡实并不能缓解浅水湖泊富营养化状况,研究结果为浅水湖泊水生植被的管理提供理论依据。     

Bloom formation of Gloeotrichia echinulata and Aphanizomenon flos-aquae in a shallow,eutrophic, Danish lake

Over a period of four years, the seasonal periodicity of dominant phytoplankton species in a shallow, eutrophic Danish lake changed markedly. Cyanophytes prevailed during the summer period of all four years. In the first three years, species of Microcystis, Anabaena and Aphanothece dominated, whereas in the fourth year of investigation, these algae were replaced by Gloeotrichia echinulata (J. E. Smith) Richter and Aphanizomenon flos-aquae (L.) Ralfs. The most striking environmental differences in the fourth year as compared with the previous three years, were an increase in tranparency, from about 0.5 meter in 1989–1991 to more than 2 metres preceding the summer maximum in 1992, and a simultaneous occurrence of low oxygen concentrations. A collapse of the fish population was followed by an increased proportion of large Cladocerans in the zooplankton. Improved light conditions at the bottom and grazing pressure from large Cladocerans favoured growth of the large colony forming blue-green algae, Gloeotrichia echinulata and Aphanizomenon flos-aquae. These species germinate from resting spores in the sediment and are able to sustain some growth there before migration to the lake water. The transfer of algal biomass from the bottom sediment to the water phase was accompanied by a marked increase in concentrations of particulate phosphorus and nitrogen in the entire lake.     

Phytoplankton community of the drinking water supply reservoir Borovitsa (South Bulgaria) with an emphasis on cyanotoxins and water quality

The phytoplankton diversity, algal biomass, and selected physicochemical parameters were investigated in the drinking water reservoir (Borovitsa) located in the Kardzhali region, Bulgaria. Particular attention was given to Cyanoprokaryota and presence of cyanotoxins in the water samples. Twenty-nine species belonging to six divisions (Cyanoprokaryota, Chlorophyta, Zygnemophyta, Dinophyta, Euglenophyta and Bacillariophyta) were identified. The microscopic examination of the phytoplankton samples showed the dominance of Ankyra judayi, Oocystis lacustris (Chlorophyta) and Aphanizomenon flos-aquae (Cyanoprokaryota) in July 2006, and Microcystis pulverea, Synechococcus elongatus (Cyanoprokaryota), Radiococcus planktonicus (Chlorophyta) and Melosira varians (Bacillariophyta) in September 2006. A blooming event due to Aphanizomenon flos-aquae was observed in July 2006. The reservoir exhibits a tendency to shift from an oligotrophic environment to a state of mesotrophy. Presence of cyanotoxins such as anatoxin-a, microcystins and saxitoxins were analyzed by HPLC and ELISA methods. Our results demonstrated the presence of anatoxin-a and microcystins (0.09 μg/L) in the raw water samples from July 2006, and saxitoxins (2.5 μg/L) and microcystins (0.18 μg/L) in the raw water samples from September 2006. The study underlines that permanent monitoring programs of Cyanoprokaryota in the reservoirs used as sources of drinking water and toxicity assessments should be implemented. Indirect exposure and transfer of cyanotoxins through food chains must also be considered.     

Notes on the growth and sporulation of a natural population of Aphanizomenon flos-aquae

Aphanizomenon flos-aquae contributed regularly to the summer phytoplankton in Kinnego Bay, a highly eutrophic part of Lough Neagh. Growth of the alga was variable in different years, and was apparently related to the duration of depletion of dissolved nitrate-nitrogen in the water. Although four growth pulses occurred during the study period, sporulation was only observed during one of these, when the population was very dense although evenly distributed down the water column. It seems that Aphanizomenon overwintered in Kinnego Bay as vegetative filaments and that production of akinetes was not necessary for perennation of the species. The observed sporulation in 1973 may have been induced by the high pH values caused by dense phytoplankton crops; no other measured environmental factors were particularly unfavourable to growth of Aphanizomenon at this time.     

Investigations of the temporal variation of cyanobacterial and other phytoplanktonic cells at the offtake of a large reservoir,and their survival following passage through it

The survival and subsequent growth potential of Anabaena spp. and other filamentous cyanobacteria and the cells of Aulacoseira spp. (diatom) and Ceratium hirundinella (dinoflagellate) following passage through the Multi Level Inlet Tower (MLIT) and offtake works at Chaffey Reservoir in New South Wales, Australia was investigated in late summer. The study aimed to test whether the phytoplankton cells were destroyed or otherwise rendered less viable during passage through the outlet works. The reservoir was strongly thermally stratified with a shallow surface mixed layer, which contributed to considerable temporal variability in the numbers of phytoplankton cells present immediately opposite the intake portal of the outlet works. To compensate, considerable replicate sampling was undertaken both upstream and downstream of the MLIT. Results indicate limited destruction of cyanobacteria, with fewer cells present immediately downstream compared to upstream. Greater destruction of cells was indicated at lower mean daily discharge rates compared to higher discharge rates. Filament lengths of both cyanobacteria and Aulacoseira were also reduced during passage. There was no apparent reduction in Ceratium cell number. Laboratory incubation studies on surviving cells collected downstream indicated no impairment on the viability of any taxa. Calculations of rates-of-strain likely to be experienced by the phytoplankton as they transited through the offtake revealed very high stress being applied to the filaments and cells at the valve, and within the spillway sections of the works. These were several orders of magnitude greater than published values shown to disrupt cells and filaments, and to impair viability for subsequent growth in laboratory studies. However, exposure times to the high rates-of-strain at Chaffey Reservoir were brief, which may reduce the impacts of the high turbulence. The conclusions were that unless cyanobacterial cell destruction during passage through an outlet works can be shown to be more effective at larger reservoirs, the withdrawal of warm, cyanobacterial infested waters from close to the surface is unlikely to provide an acceptable management action for the prevention of cold water pollution downstream. Handling editor: D. Ryder     

Importance of interspecific competition in the abundance of Aphanizomenon flos-aquae (Cyanophyceae)

The population dynamics of Aphanizomenon flos-aquae var. klebahnii Elenk. (Cyanophyceae) was studied in an artificial pond for 32 months from May 2002 to December 2004. Our previous in vitro study showed that the lower limits of water temperature and pH for its growth are within the ranges of 11°–14°C and 7.1–7.4, respectively, and it appeared that these findings are applicable to the emergence and disappearance of Ap. flos-aquae in the pond. Based on the change in the water temperature, the emergence of Ap. flos-aquae in 2004 was expected to be in late April, whereas emergence occurred after a 1-month lag period. During this period, Ankistrodesmus falcatus (Corda) Ralfs (Chlorophyceae) dominated the phytoplankton assemblage, which raised the possibility that the growth of Ap. flos-aquae was restricted by the existence of An. falcatus. We conducted mixed cultures of Ap. flos-aquae and An. falcatus at four temperatures (14°, 19°, 24°, and 29°C). After 18 days of incubation, An. falcatus dominated at 14° and 29°C whereas Ap. flos-aquae dominated at 19°C. This result indicates that a slightly higher water temperature than the growth threshold value is needed for Ap. flosaquae to outcompete An. falcatus, which agrees with the field observation. Contrary to the results of the mixed culture, the summer phytoplankton assemblage was dominated by Ap. flos-aquae, and the population of An. falcatus was less or almost absent. This variation seemed to be partly caused by the difference in nutrient conditions; concentrations of dissolved inorganic nitrogen and phosphorus in the pond were far lower than those in the culture medium. The lack of nitrogen fixation of An. falcatus seemed to be a growth disadvantage during the summer when the concentration of dissolved inorganic nitrogen was low.     

Lysis of Aphanizomenon flos-aquae (Cyanobacterium) by a bacterium Bacillus cereus

A phytoplankton-lytic (PL) bacterium, Bacillus cereus, capable of lysing the bloom-forming cyanobacterium Aphanizomenon flos-aquae was isolated from Lake Dianchi of Yunnan province, China. This bacterium showed lytic activities against a wide range of cyanobacteria/algae, including A. flos-aquae, Microcystis viridis, Microcystis wesenbergi, Microcystis aeruginosa, Chlorella ellipsoidea, Oscillatoria tenuis, Nostoc punctiforme, Anabaena flos-aquae, Spirulina maxima, and Selenastrum capricornutum. Chlorophyll a contents, phycocyanin contents, and photosynthetic activities of the A. flos-aquae decreased evidently in an infected culture for a period. Bacterium B. cereus attacked rapidly A. flos-aquae cells by cell-to-cell contact mechanism. It was shown that the lysis of A. flos-aquae began with the breach of the cyanobacterial cell wall, and the cyanobacterial cell appeared abnormal in the presence of the PL bacterium. Moreover, transmission electron microscope examinations revealed that a close contact between the bacterium and the cyanobacterium was necessary for lysis. Some slime extrusions produced from B. cereus assisted the bacterial cells to be in close association with and lyse the cyanobacterial cells. These findings suggested that this bacterium could play an important role in controlling the Aphanizomenon blooms in freshwaters.     

The distributions of autumn picoplankton in relation to environmental factors in the reservoirs along the Wujiang River in Guizhou Province,SW China

Autumn picoplankton (Synechococcus, picoeukaryotes and heterotrophic bacteria) and environmental factors have been investigated in a series of reservoirs along the Wujiang River in Guizhou Province, SW China. The average abundances of Synechococcus, picoeukaryotes and heterotrophic bacteria was 10⁴, 10² and 10⁶ cells ml⁻¹, respectively. In autumn meso-eutrophic reservoirs, thermal stratification was clear and abundances of different picoplankton groups in release water was low; whereas these phenomena were not obvious in autumn hypereutrophic reservoir. Picoplankton numbers decreased with increasing water depth and showed a positive correlation with water temperature, which reflected the importance of light and temperature on the picoplankton growth. Contribution of Synechococcus to total phytoplankton production and contribution of picoeukaryotes to total phytoplankton production asynchronous changed with varying trophic states. Synechococcus preferred meso-eutrophic reservoirs over hypereutrophic reservoir and picoeukaryotes showed no preference for the investigated reservoirs in autumn. Handling editor: L. Naselli-Flores     

Effect of Filter-feeding Zooplankton on Phytoplankton in Fish Ponds

The effect of filter-feeding zooplankton on phytoplankton was investigated in two fish ponds near Blatná (southwestern Bohemia, Czechoslovakia) using enclosures of 50 liters volume. Filter-feeding activity had a positive effect on the growth of individual net phytoplankton (> 40 μm) species except for Aphanizomenon flos-aquae and Melosira granulata. Except for Schroederia setigera nanoplankton species (< 40 μm) were suppressed by the filter-feeding zooplankton. In bags without zooplankton the addition of nutrients induced growth of nanophytoplankton species, but no significant differences were noted in bags with zooplankton.     

Effects of an Algicidal Bacterium <Emphasis Type="Italic">Pseudomonas mendocina</Emphasis> on the Growth and Antioxidant System of <Emphasis Type="Italic">Aphanizomenon flos-aquae</Emphasis>

Evident effect of an algicidal bacterium Pseudomonas mendocina on the growth and antioxidant system of Aphanizomenon flos-aquae was detected in this experiment. Seven parameters including the chlorophyll a contents, Fv/Fm values, reactive oxygen species (ROS), malonaldehyde (MDA), catalase (CAT), peroxide dismutase (POD), and superoxide dismutase (SOD) were tested in the cyanobacterium A. flos-aquae cells after inoculation with the algicidal bacterium Pseudomonas mendocina DC10. It was shown from the experiment that the growth of the treated cyanobacterium A. flos-aquae was significantly restrained, which was expressed as great reductions in the chlorophyll a contents and Fv/Fm values. At the same time, the treated cyanobacterial cells exhibited an obvious increase in the production of ROS and MDA compared with the control. CAT and POD activities in the treated group kept at high level, however, they both reduced significantly on day 6. SOD activities in the treated A. flos-aquae showed obvious declines after inoculation, and great augmentations on day 3 and 4, thereafter, they kept in a declining tendency. The results showed the oxidative stresses induced by the bacterium could be a killing agent of the cyanobacterium A. flos-aquae cells.     

Cyanobacterial blooms and water quality in Greek waterbodies

The cyanobacterial species composition of nine Greek waterbodies of different type and trophic status was examined during the warm period of the year (May–October). Cyanobacterial water blooms were observed in all waterbodies. Forty-six cyanobacterial taxa were identified, 11 of which are known to be toxic. Eighteen species are reported for the first time in these waterbodies, 8 of which are known to produce toxins. Toxin producing species were found in all of the waterbodies and were primarily dominant in bloom formations (e.g., Microcystis aeruginosa, Anabaena flos-aquae, Aphanizomenon flos-aquae and Cylindrospermopsis raciborskii). Cosmopolitan species (e.g., M. aeruginosa), pantropic (e.g., Anabaenopsis tanganyikae) and holarctic species (e.g., Anabaena flos-aquae) were encountered. Shallow, eutrophic waterbodies had blooms dominated by Microcystis species and were characterized by phytoplankton association M. Anabaena and Aphanizomenon species of association H were dominant in waterbodies with low dissolved inorganic nitrogen and thermal stratification in the summer. Total cyanobacterial biovolumes (CBV) ranged from 7 to 9,507 cm³ m⁻³ and were higher than Alert Level 2 and Guidance Level 2 (10 cm³ m⁻³; World Health Organization; WHO) in seven of the waterbodies. Chlorophyll a concentrations ranged from 6 to 90,000 mg m⁻³ and were higher than Alert Level 2 and Guidance Level 2 (50 mg m⁻³; WHO) in eight of the waterbodies. There is also an elevated risk of acute toxicosis (Guidance Level 3; WHO) in five waterbodies. Water of an undesirable quality, hazardous to humans and animals occurs in several Greek waterbodies.     

Daily changes of uptake of inorganic carbon and nitrogen,and their relation to phytoplankton blooms in late spring-early summer in Lake Nakanuma,Japan

Daily changes of inorganic carbon and nitrogen uptake were measured in May in 1986 in Lake Nakanuma, Japan. Uptake of inorganic carbon and ammonium in the light-bottle experiments in the 1 m layers, showed daily changes similar to chlorophyll a changes, though the uptake activities peaked before chlorophyll a peaks (phytoplankton blooms) appeared. Potential growth rates of phytoplankton and observed growth rates were calculated from the uptake rates and chlorophyll a changes. The potential growth rates did not always correspond to the observed growth rates. The potential growth rates did not correlate with the loss rates. The correlation between the observed growth rates and the loss rates was better. These results suggest that though the increase of uptake activities may be necessary for occurrence of phytoplankton blooms, loss processes may affect the occurrence of blooms.     

Utilization of cyanobacteria by Brachionus calyciflorus: Anabaena flos-aquae (NRC-44-1) as a sole or complementary food source

The rotifer Brachionus calyciflorus can utilize the cyanobacterium Anabaena flos-aquae as either a sole or supplementary food source in laboratory culture. Positive population growth rates accompany food densities of 10 or 100 µg dry weight ml^–1, but slightly negative rates are found at a lower density (1.0 µg ml^–1). These results are consistent for rotifers feeding on two strains of A. flos-aquae, UTEX-1444 and NRC-44-1, with slightly enhanced survivorship and reproduction with the latter food. A 1:1 mixture (by dry weight) of Euglena gracilis and A. flos-aquae (NRC-44-1) produces survivorship comparable to that of control rotifer cohorts fed E. gracilis alone, but elicits significantly greater fecundity and population growth rates than found with the control food suspension at the same biomass density.     

Cyanophycean blooms in the reservoir of Val Joly (northern France) and their development in downstream rivers

A study was carried out in 1992 on the Val Joly reservoir and the rivers situated downstream: the River Helpe Majeure and the canalized Sambre into which it flows 40 km below the reservoir. Each year, blooms of blue-green algae occur in the reservoir. They are dominated by species such as Aphanizomenon flos-aquae (L.) Ralfs in 1991 or Pseudanabaena tenuis Koppe and Oscillatoria pseudogeminata G. Schmid in 1992. A typical fluvial composition of the phytoplankton was mostly restored above the confluence with the Sambre. Nevertheless, significant quantities of Oscillatoria and Pseudanabaena have been observed in the River Sambre, just below its confluence with the Helpe Majeure. The environmental conditions in the River Sambre do not allow Aphanizomenon to develop, but could favour Oscillatoria blooms.     

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.     

The influence of daylength,light intensity and temperature on the growth rates of planktonic blue-green algae

The in vitro growth rates under continuous light of the four dominant blue-green algae in Lough Neagh, Anabaena flos-aquae Bréb., Aphanizomenon flos-aquae Ralfs fa. gracile Lemm., Oscillatoria agardhii Gom. and Oscillatoria redekei van Goor were slower than in situ rates from Lough Neagh that had been corrected for hours of light received by the algae. However, by culturing on a 6: 18 light-dark cycle in vitro growth rates were obtained that were similar to the in situ rates. Under continuous light small species showed the fastest growth with Oscillatoria redekei the dominant species. However, this pattern was almost completely reversed under the light-dark cycle with Oscillatoria redekei only exhibiting the fastest growth rate under low light conditions. This observation showed agreement with Lough Neagh field data which showed that Oscillatoria redekei reached its maximum crop in April while the other three species were dominant during the summer months. Compared to the generally assumed high thermal tendency of blue-green algae the temperature maxima of the four species were low. No growth was observed at 35°C for any species while Anabaena flos-aquae was severely inhibited at 25°C.     

Some ecological effects of artificial circulation on a small eutrophic lake with particular emphasis on phytoplankton II. Kezar lake experiment, 1969

A small eutrophic New Hampshire lake was artificially circulated (mixed) from 28 May to 15 September, 1969, inclusive, to impede the annual bloom of a noxious blue-green alga; yet Aphanizomenon flos-aquae bloomed immediately after mixing was commenced. The bloom collapsed in early July; it was succeeded by heavy growth of predominantly chlorophycean taxa. In an in vitro experiment Aphanizomenon flos-aquae did reattain bloom proportions when the influence of artificial circulation was removed. Other phytoplankton exhibited population pulses only when the dominance of cyanophycean and chlorophycean taxa were in transition. Mixing maintained uniform vertical populations of all phytoplankton. Changes in water transparency attended fluctuations in phytoplankton abundance. An isothermic condition was maintained over the test period, which increased the lake's heat budget, and most chemical nutrients were distributed isometrically in the water column. Increased concentrations were exhibited by Ca, Cl, Cu, K, Mg, SiO₂ and Zn. Sodium was not affected by mixing. Levels of Fe, Mn, Zn, phosphate and ammonia, nitrate, and organic nitrogen were influenced by phytoplankton. Mixing could not maintain orthograde profiles of dissolved O₂ and CO₂ when dense populations of phytoplankton prevailed. Variations in chlorophyll-a followed closely changes in phytoplankton abundance. Its degradation to pheo-pigments appeared to be less for a bloom of Aphanizomenon flos-aquae than during dense growth of chlorophycean taxa. Rates of photosynthesis were considerably greater when the latter algae were predominant. Extracellular release of organic carbon usually increased with depth; it amounted to 19.4 percent of the total carbon fixed in the euphotic zone.     

Effectiveness of phytoplankton control by large-bodied and small-bodied zooplankton

Employingin situ enclosures containing inocula of the lake zooplankton (mainlyDaphnia galeata, Daphnia cucullata andBosmina spp.) from a moderately eutrophic Lake Ros (Northern Poland) or large-bodiedDaphina magna, the following observations on succession of phytoplankton were made: 1) whereasD. magna could control the density of all the photoplankton size classes, the lake zooplankton could not suppress the large-sized phytoplankters or net phytoplankton; 2) the lake zooplankton was able to control the density of small algae (< 50μm), but its effect on large algae may be opposite: a promotion of net phytoplankton growth by removing small-sized algae which can out-compete net phytoplankton for limited PO₄-P resources (<5μg P l⁻¹). Since efficiency of phytoplankton density control byD. magna decreased with an increase in net phytoplankton abundance, biomanipulation could not be successful without introducing or maintaining a high population of large-bodied cladoceran species before high densities of large algae would make the control of phytoplankton inefficient.