Understanding interactive inducible defenses of Daphnia and its phytoplankton prey

Cyanobacterial and zooplankton inducible defenses are important but understudied process that regulate the trophic interactions of freshwater ecosystem. Daphnia due to its large size is considered an important zooplankton with the high potential to control cyanobacterial blooms. It has been shown that Daphnia through maternal induction transfer tolerance to their next generation against Microcystis toxicity. Maternal induction has been investigated in different Daphnia species without considering phenotypic plasticity of prey. Laboratory experiments were performed to explore cyanobacteria-Daphnia inducible defenses in order to better understand their interactions. Two Daphnia species were fed either with Microcystis aeruginosa PCC7806 (Ma) or Microcystis flos-aquae (Mf) mixed with Chlorella vulgaris (Cv) (exposed Daphnia), and or pure Cv (unexposed Daphnia). Exposed prey cultures were produced by prior exposure to Daphnia infochemicals. Neonates produced by exposed and unexposed Daphnia were fed with mixed diet (Microcystis + Cv) of either exposed and or unexposed prey. Growth parameters and toxin production of exposed prey cultures were significantly different than that of control. Exposed Daphnia fecundity and survival was higher as compared to unexposed Daphnia. Growth and reproduction was reduced in exposed Daphnia when fed with exposed prey as compared to those fed with unexposed prey. This study provides information on the interactive inducible defenses between cyanobacteria and its grazer under laboratory conditions and may increase our understanding of cyanobacteria and Daphnia interactions in the freshwater ecosystem.     

Impact of Toxic Cyanobacterial Blooms on Eurasian Perch (Perca fluviatilis): Experimental Study and In Situ Observations in a Peri-Alpine Lake

Due to the importance of young-of-the-year (YOY) perch in the peri-alpine regions where they are consumed, the microcystin (MC) contamination of YOY perch was analysed both in field (Lake Bourget, France) and experimentally using force-feeding protocols with pure MCs. In-situ, schools of YOY perch present in the epilimnion of the lake were never found in direct contact with the P. rubescens blooms that were present in the metalimnion. However, MCs were detected in the muscles and liver of the fish and were thus assumed to reach YOY perch through dietary routes, particularly via the consumption of MC-containing Daphnia. Force-feeding experiment demonstrates the existence of MC detoxification/excretion processes and suggests that in situ, YOY perch could partly detoxify and excrete ingested MCs, thereby limiting the potential negative effects on perch populations under bloom conditions. However, because of chronic exposure these processes could not allow for the complete elimination of MCs. In both experimental and in situ studies, no histological change was observed in YOY perch, indicating that MC concentrations that occurred in Lake Bourget in 2009 were too low to cause histological damage prone to induce mortality. However, Deoxyribonucleic acid (DNA) damages were observed for both the high and low experimental MC doses, suggesting that similar effects could occur in situ and potentially result in perch population disturbance during cyanobacterial blooms. Our results indicate the presence of MCs in wild perch, the consumption of this species coming from Lake Bourget is not contested but more analyses are needed to quantify the risk.     

Response of Daphnia's Antioxidant System to Spatial Heterogeneity in Cyanobacteria Concentrations in a Lowland Reservoir

Many species and clones of Daphnia inhabit ecosystems with permanent algal blooms, and they can develop tolerance to cyanobacterial toxins. In the current study, we examined the spatial differences in the response of Daphnia longispina to the toxic Microcystis aeruginosa in a lowland eutrophic dam reservoir between June (before blooms) and September (during blooms). The reservoir showed a distinct spatial pattern in cyanobacteria abundance resulting from the wind direction: the station closest to the dam was characterised by persistently high Microcystis biomass, whereas the upstream stations had a significantly lower biomass of Microcystis. Microcystin concentrations were closely correlated with the cyanobacteria abundance (r = 0.93). The density of daphniids did not differ among the stations. The main objective of this study was to investigate how the distribution of toxic Microcystis blooms affects the antioxidant system of Daphnia. We examined catalase (CAT) activity, the level of the low molecular weight antioxidant glutathione (GSH), glutathione S-transferase (GST) activity and oxidative stress parameters, such as lipid peroxidation (LPO). We found that the higher the abundance (and toxicity) of the cyanobacteria, the lower the values of the antioxidant parameters. The CAT activity and LPO level were always significantly lower at the station with the highest M. aeruginosa biomass, which indicated the low oxidative stress of D. longispina at the site with the potentially high toxic thread. However, the low concentration of GSH and the highest activity of GST indicated the occurrence of detoxification processes at this site. These results demonstrate that daphniids that have coexisted with a high biomass of toxic cyanobacteria have effective mechanisms that protect them against the toxic effects of microcystins. We also conclude that Daphnia''s resistance capacity to Microcystis toxins may differ within an ecosystem, depending on the bloom''s spatial distribution.     

Role of Microcystins in Poisoning and Food Ingestion Inhibition of Daphnia galeata Caused by the Cyanobacterium Microcystis aeruginosa

The effects of microcystins on Daphnia galeata, a typical filter-feeding grazer in eutrophic lakes, were investigated. To do this, the microcystin-producing wild-type strain Microcystis aeruginosa PCC7806 was compared with a mcy⁻ PCC7806 mutant, which could not synthesize any variant of microcystin due to mutation of a microcystin synthetase gene. The wild-type strain was found to be poisonous to D. galeata, whereas the mcy⁻ mutant did not have any lethal effect on the animals. Both variants of PCC7806 were able to reduce the Daphnia ingestion rate. Our results suggest that microcystins are the most likely cause of the daphnid poisoning observed when wild-type strain PCC7806 is fed to the animals, but these toxins are not responsible for inhibition of the ingestion process.     

A comparison of the sensitivities of Daphnia magna and Daphnia pulex to six different cyanobacteria

Cyanobacteria have become an important environmental concern due to their ability to produce a wide range of natural toxins. At present, very few studies describe concentration response curves for cyanobacteria other than Microcystis. However, field evidence highlights that both cyanobacterial concentration as well as cyanobacterial species composition vary considerably with season and year. Therefore, the aim of this study was to investigate the effects of different cyanobacteria at various concentrations of these cyanobacteria in the diet on the reproduction of Daphnia pulex and Daphnia magna. Those two species were chosen to assess whether the cyanobacteria-daphnid dynamics could be generalized for the Daphnia genus. Results demonstrated that both slope and EC50 of the concentration response curves depend upon the Daphnia species, the cyanobacteria species and the potential interaction between the two. This has two major consequences. First, the differences in sensitivity to cyanobacteria between D. magna and D. pulex depend upon concentration of the specific cyanobacteria. Second, we noted different mechanisms of toxicity for the two zooplankton species, a more general mechanism of toxicity for D. pulex and a more specific one for D. magna. Our data therefore suggest that results of studies investigating effects of cyanobacteria at different concentrations cannot be generalized across species. Furthermore, mechanisms of toxicity are not only cyanobacteria specific, but also dependent on the exposed species, even for rather closely related species such as in the Daphnia genus. Whenever possible, we therefore propose to combine a multi-species approach together with a full concentration response analysis to reach more general conclusions concerning the effects of cyanobacteria on zooplankton.     

Food quality of mixed bacteria–algae diets for Daphnia magna

Bacteria can comprise a large fraction of seston in aquatic ecosystems and can therefore significantly contribute to diets of filter-feeding zooplankton. To assess the effect of three heterotrophic bacteria (Flavobacterium sp., Pseudomonas sp. and Escherichia coli) on survival, growth and egg production of juvenile Daphnia magna during six-day growth experiments, five ratios of bacteria–Scenedesmus obliquus mixtures were fed. Potential growth-limiting effects mediated by essential biochemicals were assessed upon supplementation of pure bacterial diets with a sterol (cholesterol) or a polyunsaturated fatty acid (EPA). Pure bacterial diets always had detrimental effects on Daphnia. However, cholesterol supplementation of Flavobacterium sp. enhanced growth rates of Daphnia. Diets containing Pseudomonas impaired Daphnia growth even at low dietary proportions (20%), indicating their toxicity. In contrast, Daphnia grew at relative high dietary proportions of Flavobacterium sp. and E. coli (80–50%). In fact, diets containing small proportions of these heterotrophic bacteria (Flavobacterium ≤50%, E. coli 20%) even significantly increased Daphnia growth rates compared to pure algal diets, indicating a nutritional upgrading by these bacteria. Our results suggest that the relative contribution of bacteria and phytoplankton to total dietary carbon as well as their phylogenetic composition strongly influence Daphnia fitness and potentially other filter-feeding zooplankton under field conditions.     

Does microcystin disrupt the induced effect of Daphnia kairomone on colony formation in Scenedesmus?

In natural aquatic system, Scenedesmus and Microcystis species usually coexist. Microcystins are released into water after lysis of Microcystis cells during the collapse of heavy blooms. The released toxins can then come into contact with a wide range of aquatic organisms. In this study, we used filtered Daphnia test water containing kairomone from Daphnia magna to stimulate the inducible colony formation in Scenedesmus obliquus under microcystin-contaminated system, to examine how microcystin affects the induced effect of Daphnia kairomone on colony formation in S. obliquus. The results showed neither microcystin nor Daphnia kairomone affected the growth of S. obliquus. Microcystin neither promoted nor impaired the overall Daphnia-induced colony formation in S. obliquus, except reducing the proportion of eight-celled colonies on day 2, indicating that the effect of microcystin was just short-term and in general did not disrupt grazer-induced colony formation of S. obliquus.     

The effects of a microcystin-producing and lacking strain of Microcystis on the survival of a widespread tropical copepod (Notodiaptomus iheringi)

The tropical copepod Notodiaptomus iheringi (N. iheringi) is an ideal subject for studying zooplankton responses to cyanobacteria because it co-exists with permanent blooms across widespread regions in South America in high abundance. Single and mixed diets containing Cryptomonas and either a microcystin-producing (MC+) or microcystin-lacking (MC?) Microcystis were offered to N. iheringi at different proportions in a 10-day laboratory survival test to distinguish between the effects of toxicity versus nutrition. As expected, the pure MC+ Microcystis diet caused acute toxicity, indicated by high mortality compared to starved copepods. Both Microcystis strains were ingested in a 3-h short-term grazing experiment with pure diets. Despite its toxicity as the sole food source, survival was unaffected by MC+ Microcystis in mixed food diets. Even when MC+ Microcystis was 90% of the total food, survival was similar to the control with 10% Cryptomonas only. Hence, the survival in mixed food diets was controlled by the amount of Cryptomonas, not Microcystis. Previous reports show strong negative effects of Microcystis on copepod survival despite abundant high-quality food. Although this is the first example of copepods avoiding acute Microcystis toxicity in mixed diets, it could be a common trait where permanent blooms dominate the ecosystem.     

Midsummer decline of a Daphnia population attributed in part to cyanobacterial capsule production

A midsummer decline in a Daphnia population has been documentedin Lake Winnebago, Wisconsin, USA, similar to that which typicallyoccurs in other freshwater lakes throughout the world. We investigatedthe role of two products of the cyanobacterium Microcystis aeruginosa(microcystin and polysaccharide capsular matrix) in contributingto this population decrease by utilizing laboratory and lakestudies. Feeding on either the microcystin-producing, minimallyencapsulated M. aeruginosa strain PCC 7820 or the highly encapsulated,non-microcystin-producing strain C3-40, resulted in decreasedsurvival when compared with animals fed chlorophytes. The survivorshipof Daphnia fed C3-40 cells washed to remove the capsule increasedby 30% over that seen in animals fed encapsulated C3-40. Feedingpurified microcystin or capsule alone decreased Daphnia survivalto a greater degree than did starvation. Physiological studiesconducted while Daphnia were exposed to these food sources showedthat ingestion of capsular material resulted in increased post-abdominalclaw movements and decreased mandibular movements. Concurrently,elevated respiration rates were measured in Daphnia, implyingthat capsule increased the energy expended by these animalsthrough increased attempts to reject the material and decreasedfood intake. Lake studies reflected the results of the laboratoryexperiments. The midsummer decline of the Daphnia populationoccurs as the Microcystis biovolume increases and both microcystinand capsular matrix levels rise. While both cyanobacterial productsmay contribute to the midsummer decline in Daphnia pulicaria,laboratory studies suggest that encapsulation may play the greaterrole.     

Toxin production of cyanobacteria is increased by exposure to zooplankton

1. Cyanobacterial toxin production in response to direct and indirect zooplankton feeding activity was examined using four strains of Microcystis aeruginosa, of which three were previously reported to be toxic to zooplankton and one non‐toxic. Direct (Microcystis cultured with zooplankton) and indirect effects (Microcystis cultured with filtered zooplankton culture media, ZCMF) were tested for the zooplankton species, Moina macrocopa, Daphnia magna or D. pulex. 2. With direct exposure to zooplankton, increased mass‐specific microcystin productions occurred in all Microcystis strains, with mean microcystin concentrations up to five times greater (61.5–177.3 μg g^?1 dry cell) than the controls. 3. With indirect exposure, mass‐specific microcystin production increased over controls in three strains of M. aeruginosa. Mean maximum concentrations of microcystin during the experiment were 92.6–125.7 μg g^?1 dry cell. 4. These results suggest that several strains of Microcystis aeruginosa increased toxin production in response to direct and indirect exposure to herbivorous zooplankton of several species, and support the hypothesis that this response is an induced defence mediated by the release of info‐chemicals from zooplankton.     

Feeding behaviour,efficiency and food preference in yabbies <Emphasis Type="Italic">Cherax</Emphasis><Emphasis Type="Italic"> destructor</Emphasis>

The influence of body size on the consumption of live zooplankton (Daphnia spp.) by freshwater crayfish was examined using yabbies (Cherax destructor) ranging from 5 to 45 g. Food preference between live zooplankton and inert pellets was also assessed under experimental conditions. In experimental tanks, yabbies of four size classes (<15, 15–24.9, 25–34.9 and 35–45 g) were presented with live Daphnia. All yabbies were held in separate tanks with five animals per size class. In yabbies less than 15 g, the feeding mode on zooplankton involved rapid searching and probing with the first two pairs of walking legs. Once a prey was located, the chelae on the end of these walking legs would grasp the zooplankton and then rapidly move it towards the mouthparts. Yabbies larger than 25 g tended to use their walking legs to push the Daphnia nearer to their third maxilliped which would then force or scoop the zooplankton towards the mouthparts. A short-term feeding trial showed that there was no significant difference between size classes in regards to zooplankton consumption (P > 0.05). Capture efficiency of live Daphnia by yabbies less than 15 g was significantly lower (76%, P = 0.008) than the three larger size classes (93.6%). Yabbies less than 15 g consumed a significantly (P < 0.001) higher percentage (5.2%) of their body weight than the other size classes (1.1%, 0.8%, and 0.6%, respectively). In the presence of both live zooplankton and a pellet diet, yabbies spent significantly (P = 0.005) more time feeding on zooplankton (85%) than on inert pellets (15%). This was the first study to quantify zooplankton consumption by yabbies and the results provide insights into understanding the trophic role of freshwater crayfish in structuring zooplankton communities and the husbandry management of crayfish farming. Handling editor: S. I. Dodson     

Low fish predation pressure in London reservoirs: II. Consequences to zooplankton community structure

The 1992 survey of zooplankton structure in fourteen London supply reservoirs showed the overall dominance of large-bodied zooplankton, mainly species of Daphnia. These reservoirs can be considered as ‘anti-fish’ by virtue of their steeply sloping concrete or brick sides. The average biomass of large Daphnia spp (retained on a 710 μm sieve) in the total zooplankton biomass was higher than 20% for twelve out of fourteen reservoirs. The cladoceran-copepod ratio was inversely correlated with both dominance of large-bodied Daphnia magna and cladoceran body-size structure. Parallelly, there were tendency of more efficient utilization of lowered algal crops in reservoirs dominated by large-bodied Daphnia spp. A graphical model is presented which relates daphnid species composition and zooplankton size structure to a presumed gradient of fish biomass in these reservoirs.     

Context dependency of infectious disease: the cyanobacterium Microcystis aeruginosa decreases white bacterial disease in Daphnia magna

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Foraging strategies and growth inhibition in five daphnids feeding on mixtures of a toxic cyanobacterium and a green alga

1. Laboratory experiments were used to study the feeding, growth and reproduction of five daphnids in mixtures of a toxic cyanobacterium, Microcystis aeruginosa, and a green alga, Scenedesmus acutus. The mixtures included 0%, 20%, 50%, 80% and 100% Microcystis with a total food concentration of 0.5 mg C L^??1 in each treatment. The feeding rate was measured after 1 and 24 h of acclimatization to the mixtures. 2. Toxic Microcystis inhibited feeding in all the species, but they exhibited an unexpected diversity and complexity in the pattern of feeding inhibition. Daphnia magna exhibited the strongest inhibition of feeding after 1 h of exposure to toxic food, but had substantially recovered after 24 h in the same mixtures. This pattern of inhibition and recovery may balance the benefits of reduced ingestion of toxin with the disadvantage of a reduced energy intake. 3. All five daphnids grew quickly in the Scenedesmus control, whereas growth and reproduction declined with an increasing proportion of the toxic alga in the diet. Daphnia pulicaria showed the least inhibition of growth and reproduction, D. pulex showed the strongest inhibition and the three remaining species exhibited intermediate sensitivity. 4. Estimates of gross growth efficiency (GGE; growth/ingestion) provided a means for discriminating between the effects of feeding inhibition and direct toxicity on zooplankton growth. Daphnia pulex exhibited a sharp decline in GGE, suggesting that growth inhibition was a result of both feeding inhibition and direct toxicity. In contrast, D. magna exhibited a nearly constant GGE, indicating that feeding inhibition accounted for its decline in growth. However, two Daphnia species (i.e. D. pulicaria and D. galeata) exhibited improbable increases in GGE with toxic cyanobacteria, suggesting that their feeding rates were underestimated. Growth assays with sensitive and resistant zooplankton species are proposed for testing the potential impacts toxic cyanobacteria in lakes.     

Daphnia community analysis in shallow Kenyan lakes and ponds using dormant eggs in surface sediments

1. Water fleas of the genus Daphnia are considered rare in tropical regions, and information on species distribution and community ecology is scarce and anecdotal. This study presents the results of a survey of Daphnia species distribution and community composition in 40 standing waterbodies in southern Kenya. The study sites cover a wide range of tropical standing aquatic habitats, from small ephemeral pools to large permanent lakes between approximately 700 and 2800 m a.s.l. Our analysis combines data on Daphnia distribution and abundance from zooplankton samples and dormant eggs in surface sediments. 2. Nearly 70% (27 of 40) of the sampled waterbodies were inhabited by Daphnia. Although their abundance in the active community was often very low, this high incidence shows that Daphnia can be equally widespread in tropical regions as in temperate regions. 3. Analysis of local species assemblages from dormant eggs in surface sediments was more productive than snapshot sampling of zooplankton communities. Surface‐sediment samples yielded eight Daphnia species in total, and allowed the detection of Daphnia in 25 waterbodies; zooplankton samples revealed the presence of only four Daphnia species in 16 waterbodies. 4. Daphnia barbata, D. laevis, and D. pulex were the most frequently recorded and most abundant Daphnia species. Canonical correspondence analysis of species–environment relationships indicates that variation in the Daphnia community composition of Kenyan waters was best explained by fish presence, temperature, macrophyte cover and altitude. Daphnia barbata and D. pulex tended to co‐occur with each other and with fish. Both species tended to occur in relatively large (>10 ha) and deeper (>2 m) alkaline waters (pH 8.5). Daphnia laevis mainly occurred in cool and clear, macrophyte‐dominated lakes at high altitudes.     

Effects of Cell-Bound Microcystins on Survival and Feeding of Daphnia spp.

The influence of cell-bound microcystins on the survival time and feeding rates of six Daphnia clones belonging to five common species was studied. To do this, the effects of the microcystin-producing Microcystis strain PCC7806 and its mutant, which has been genetically engineered to knock out microcystin synthesis, were compared. Additionally, the relationship between microcystin ingestion rate by the Daphnia clones and Daphnia survival time was analyzed. Microcystins ingested with Microcystis cells were poisonous to all Daphnia clones tested. The median survival time of the animals was closely correlated to their microcystin ingestion rate. It was therefore suggested that differences in survival among Daphnia clones were due to variations in microcystin intake rather than due to differences in susceptibility to the toxins. The correlation between median survival time and microcystin ingestion rate could be described by a reciprocal power function. Feeding experiments showed that, independent of the occurrence of microcystins, cells of wild-type PCC7806 and its mutant are able to inhibit the feeding activity of Daphnia. Both variants of PCC7806 were thus ingested at low rates. In summary, our findings strongly suggest that (i) sensitivity to the toxic effect of cell-bound microcystins is typical for Daphnia spp., (ii) Daphnia spp. and clones may have a comparable sensitivity to microcystins ingested with food particles, (iii) Daphnia spp. may be unable to distinguish between microcystin-producing and -lacking cells, and (iv) the strength of the toxic effect can be predicted from the microcystin ingestion rate of the animals.     

The role of microcystins in maintaining colonies of bloom-forming Microcystis spp

Microcystis is a cosmopolitan genus of cyanobacteria and occurs in many different forms. Large surface blooms of the cyanobacterium are well known in eutrophic lakes throughout the globe. We evaluated the role of microcystins (MCs) in promoting and maintaining bloom-forming cell aggregates at environmentally relevant MC concentrations (0.25-10 μg l(-1)). MCs significantly enhanced Microcystis colony sizes. Colonial diameters in microcystin-RR (MC-RR)-treated cultures (at 1 μg l(-1)) were significantly larger than control colonies, by factors of 1.5, 2.6 and 2.7 in Microcystis wesenbergii DC-M1, M. ichthyoblabe TH-M1 and Microcystis sp. FACHB1027 respectively. Depletion of extracellular MC concentrations caused Microcystis colony size to decrease, suggesting that released MCs are intimately involved in the maintenance of Microcystis colonial size. MC-RR exposure did not influence Microcystis growth rate, but did significantly increase the production of extracellular polysaccharides (EPS). In addition, MC-RR exposure appeared to trigger upregulation of certain parts of four polysaccharide biosynthesis-related genes: capD, csaB, tagH and epsL. These results strongly indicate that induction of polysaccharides by MC-RR was the major mechanism through which MCs enhanced colony formation in Microcystis spp. Cellular release of MCs, therefore, may play a key role in the persistence of algal colonies and the dominance of Microcystis.     

Cadmium Toxicity to Microcystis aeruginosa PCC 7806 and Its Microcystin-Lacking Mutant

The adverse effects of microcystin (MC) produced by cyanobacteria have drawn considerable attention from the public. Yet it remains unclear whether MC confers any benefits to the cyanobacteria themselves. One suggested function of MC is complexation, which may influence the bioaccumulation and toxicity of trace metals. To test this hypothesis, we examined Cd toxicity to wild-type Microcystis aeruginosa PCC 7806 (WT) and its MC-lacking mutant (MT) under nutrient-enriched (+NP), phosphorus-limited (-P), and nitrogen-limited (-N) conditions. The accumulation of Cd and the biochemical parameters associated with its detoxification [total phosphorus (TP), inorganic polyphosphate (Poly-P), and glutathione (GSH) in the cells as well as intra- and extra-cellular carbohydrates] were quantified. Although the –P cyanobacteria accumulated less Cd than their +NP and –N counterparts, the different nutrient-conditioned cyanobacteria were similarly inhibited by similar free ion concentration of Cd in the medium ([Cd²⁺]_F). Such good toxicity predictability of [Cd²⁺]_F was ascribed to the synchronous decrease in the intracellular concentrations of Cd and TP. Nevertheless, Cd toxicity was still determined by the intracellular Cd to phosphorus ratio (Cd/P), in accordance with what has been reported in the literature. On the other hand, the concentrations of TP, Poly-P, and carbohydrates went up, but GSH concentration dropped down with the enhancement of [Cd²⁺]_F, indicating their association with Cd detoxification. Although the inactivation of MC peptide synthetase gene had some nutrient and Cd concentration dependent effects on the parameters above, both cyanobacterial strains showed the same Cd accumulation ability and displayed similar Cd sensitivity. These results suggest that MC cannot affect metal toxicity either by regulating metal accumulation or by altering the detoxification ability of the cyanobacteria. Other possible functions of MC need to be further investigated.     

Planktonic food chains of a highly humic lake

The development and metabolism of epilimnetic plankton from a highly humic lake was followed in late summer, when the predominant zooplankton species, Daphnia longispina, was very abundant (ca. 200 ind. l^?1). The experiment was made in two tanks: one with an unaltered plankton assemblage and one with larger zooplankton removed. The scarce phytoplankton community was also simple, consisting mainly of one Cryptomonas and two Mallomonas species. The abundance and species composition of smaller plankton was heavily influenced by grazing of Daphnia. In particular, the biomass, of heterotrophic flagellates increased after the removal of Daphnia. The biomass and production of bacterioplankton were not affected, and remained several times higher than that of phytoplankton. Bacterial production and grazing on bacteria were balanced, and when Daphnia was removed its grazing activity was compensated by flagellates. The removal of Daphnia did not affect the respiration or community net production of plankton. Among organisms smaller than zooplankton, bacteria seemed to be responsible for most of the respiration. The community net production was consistently negative even at the water surface, indicating an allochthonous carbon source. The results suggest that phytoplankton primary production was insufficient for the secondary production in the epilimnetic water of the study lake. The food requirements of bacteria and zooplankton, as well as of flagellates, each exceeded that supplied by phytoplankton primary production. The simple food chains in this experiment made it possible to reveal the functioning of the community so completely that dissolved organic matter is certainly comparable to or exceeds the importance of phytoplankton primary production as an energy and carbon source for food webs in this humic lake.     

The abundance of microcystin-producing genotypes correlates positively with colony size in Microcystis sp. and determines its microcystin net production in Lake Wannsee

The working hypotheses tested on a natural population of Microcystis sp. in Lake Wannsee (Berlin, Germany) were that (i) the varying abundance of microcystin-producing genotypes versus non-microcystin-producing genotypes is a key factor for microcystin net production and (ii) the occurrence of a gene for microcystin net production is related to colony morphology, particularly colony size. To test these hypotheses, samples were fractionated by colony size with a sieving procedure during the summer of 2000. Each colony size class was analyzed for cell numbers, the proportion of microcystin-producing genotypes, and microcystin concentrations. The smallest size class of Microcystis colonies (<50 microm) showed the lowest proportion of microcystin-producing genotypes, the highest proportion of non-microcystin-producing cells, and the lowest microcystin cell quotas (sum of microcystins RR, YR, LR, and WR). In contrast, the larger size classes of Microcystis colonies (>100 microm) showed the highest proportion of microcystin-producing genotypes, the lowest proportion of non-microcystin-producing cells, and the highest microcystin cell quotas. The microcystin net production rate was nearly one to one positively related to the population growth rate for the larger colony size classes (>100 microm); however, no relationship could be found for the smaller size classes. It was concluded that the variations found in microcystin net production between colony size classes are chiefly due to differences in genotype composition and that the microcystin net production in the lake is mainly influenced by the abundance of the larger (>100- microm) microcystin-producing colonies.