Life history consequences for Daphnia pulex feeding on nutrient-limited phytoplankton

1. The growth and feeding of Daphnia pulex De Geer on different algal species was examined. The green algae Chlamydomonas reinhardtii Dangeard and Scenedesmus acutus Meyen, the diatom Synedra tenuissima Kützing, the cryptophyte Cryptomonas pyrenoidifera Geitler and the cyanobacterium Microcystis aeruginosa Kützing were cultured in non-limiting and in N- or P-limiting medium and used as food for D. pulex.
2. Growth limitations were reflected in the elemental and biochemical composition and the morphological characteristics of the algal resources.
3. The clearance rates of D. pulex feeding on nutrient-limited algae were reduced. This was not observed when nutrient-limited mutant Chlamydomonas cells without cell walls were used as food, indicating that the cell wall may play an important part.
4. In comparison with animals grown on nutrient-sufficient cells, nutrient-limited algae resulted in smaller body length, reduced brood sizes, reduced size at maturity, increased age at first reproduction and, consequently, in reduced Daphnia population growth rates.
5. Daphnia population growth rates ( r ) were negatively correlated with the C : P ratio and the carbohydrate content of the food. Moreover, significant correlations between r and clearance rates were found.
6. The observed differences in the grazing and the life history parameters of Daphnia feeding on non-limited and nutrient-limited algae may be the result of both reduced nutritional value and reduced digestibility of nutrient-limited algae.     

UV-induced changes in phytoplankton cells and its effects on grazers

1. This review addresses the effects of UV-radiation on the morphology and biochemistry of phytoplankton and the potential effects on grazers.
2. UVA and UVB radiation inhibit the uptake of inorganic nutrients in phytoplankton. Reduced rates of ammonium and nitrate uptake in marine diatoms, and reduced uptake of phosphorus in freshwater flagellates are reported. The effects on cell stoichiometry are not settled.
3. UVA and UVB radiation promote increased cell volumes owing to a decoupling between the photosynthetic processes and cell division. Loss or inactivation of flagellae and loss of motility are also reported for a number of phytoplankton species.
4. UVA and UVB radiation may affect major biochemical constituents. Accumulation of intracellular, photosynthetic products (lipids or carbohydrates) is a common, although not unique, property of UV-stressed algae. Fatty acid (FA) profiles seem susceptible to UV radiation. A relative increase of short-chained, and a decrease in polyunsaturated FA (PUFA) are reported. The important membrane FA like eicosapentaenoic acid (EPA, 20 : 5ω3) and docosahexaenoic acid (DHA, 22 : 6ω3) seem particularly susceptible, owing to lipid peroxidation or reduced biosynthesis.
5. UV-related responses are highly dependent on taxonomy, cell-cycle stage, nutrient-limitation and the UV : PAR (photosynthetic active radiation)-ratio.
6. Nutrient deficiency, cell size, cell wall properties and FA can all have significant impacts on grazers. Thus the reported effects on cell morphology and biochemical constituents could have profound effects on grazers and energy transfer in aquatic foodwebs.     

The effect of fungal parasitism on the succession of diatoms in Lake Maarsseveen I (The Netherlands)

SUMMARY. During late winter and early spring, diatoms are predominant in Lake Maarsseveen I and show a succession in blooming. To elucidate the role of parasitism in this succession, samples were taken once a week in 1978, 1979and 1980 and examined for fungal parasites, further, factors which presumably influenced the phytoplankton growth. including temperature and chemical compounds, were studied. Parasites were found on Asterionella formosa Hass., Stephanodiscus astraea (Ehr) Grun. and Cyclotella comta (Ehr) Kütz. The chytrid fungus Zygorhizidium planktonicum
Canter was parasitic tin Asterionella formosa. This was the only case where high infection percentages were registered. The developmental phases of Zygorhizidium plunktonicum on Asterionella formosa were studied. The fungus was capable of affecting the spring increase and bringing about a premature end to the bloom. Only when the fungus was temporarily inhibited in its parasitic activity (this phenomenon coincided with periods of frost) was A. formosa able to reach a high abundance. Severe parasitism on A. formosa favoured the development of other diatoms (mainly of Fragilaria crotoriensis Kitt., Stephanodiscus hantzschii Grun. and Stephanodiscus astraea) .
Host-parasite interrelationship studies performed in the laboratory showed that temperature can be an important environmental factor associated with epidemics of Zygorhizidium plankiomcum on Asterionella formosa. At very low temepratures (1.5. 1) the fungal activity was inhibited while Asterionella fortnosa still grew well. However, at 5, 10 and 18 the fungus manifested a high infection rate and was able to overtake A sterionella .     

Allelopathic effect of the aquatic macrophyte, Stratiotes aloides, on natural phytoplankton

1. A survey of different Dutch Stratiotes stands showed that the density of phytoplankton (except cyanobacteria) was always higher outside S. aloides than between the rosettes of S. aloides. Analyses of water samples revealed that nutrient limitation was unlikely to have caused the lower phytoplankton biomass in the vicinity of S. aloides. 2. An in situ incubation experiment in the Danube Delta, Romania, indicated allelopathic activity against phytoplankton in S. aloides stands. The growth rate of natural phytoplankton populations exposed to water from S. aloides stands was significantly lower than that of populations that had not been in contact with S. aloides exudates. 3. A laboratory microcosm experiment showed a significantly lower phytoplankton biomass in treatments with S. aloides exudates. Nutrient concentrations and the light intensity were high enough that the lower phytoplankton biomass could not be explained by nutrient or light limitation.     

Effects of UVB-irradiated algae on life history traits of Daphnia pulex

1. The impact of ultraviolet-B (UVB)-irradiated phytoplankton on the life history parameters of Daphnia was studied. Three species of Chlorophyceae ( Chlamydomonas reinhardtii, Scenedesmus acutus and S. subspicatus ) and one species of Cryptophyceae ( Cryptomonas pyrenoidifera ) were cultured with and without exposure to UVB radiation (biologically effective dose of 3 kJ m^–2 day^–1 DNA weighted) and fed to Daphnia pulex.
2. The daphnids feeding on UVB-irradiated algae tended to be smaller in size, to produce less and smaller offspring, and to have a reduced population growth rate.
3. Fatty acid (FA) analysis of the phytoplankton showed a reduced total FA content under UVB radiation for all four species. Changes in the content of unsaturated fatty acids (UFA) were species specific.
4. The results indicate that the energy transfer between the primary producers' and grazers' level might be negatively influenced by UVB radiation. Direct effects on food quality are modest but detectable, changes in quantity or species composition may be more important in the energy transfer.     

Changes in food quality of the green alga Scenedesmus induced by Daphnia infochemicals: biochemical composition and morphology

1. The effects of Daphnia infochemicals on the morphology and biochemical composition of Scenedesmus were studied and subsequently their influence on Daphnia life history. Three species of Scenedesmus ( S. acutus, S. obliquus and S. subspicatus ) were tested for Daphnia -induced colony formation. Life history experiments were performed with Daphnia cucullata and D. magna feeding on unicellular or colonial S. acutus.
2. Colony formation was promoted when S. acutus and S. obliquus were exposed to filtered water from a D. magna culture. S. subspicatus did not form colonies when exposed to culture water, but showed some colony formation when exposed to live D. magna.
3. No clear differences were found in total lipid, protein or carbohydrate content between the unicellular and colonial ecomorphs of the three Scenedesmus species. However, fatty acid (FA) concentration and composition were changed. Total FA as a percentage of frozen dry weight (DW) were increased in colonies. The ratio of monounsaturated (MUFA) and polyunsaturated fatty acids (PUFA) and the ratio of ω3 : ω6 fatty acids decreased in colonies.
4. A short-term grazing experiment with three size classes of Scenedesmus revealed lower clearance rates for small Daphnia when feeding on large colonial Scenedesmus.
5. Life history parameters of both Daphnia species were influenced by food type. The intrinsic rate of population increase ( r) was significantly lower for daphnids feeding on colonies.
6. The negative influence of colonial Scenedesmus on population growth of Daphnia seems to be determined by algal morphology rather than biochemical composition.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

The effect of a mixotrophic chrysophyte on toxic and colony-forming cyanobacteria

1. In order to test the effect of Ochromonas sp. , a mixotrophic chrysophyte, on cyanobacteria, grazing experiments were performed under controlled conditions. We studied grazing on three Microcystis aeruginosa strains, varying in toxicity and morphology, as well as on one filamentous cyanobacterium, Pseudanabaena sp. Furthermore, we analysed the co-occurrence of Ochromonas and Microcystis in natural systems in relation to various environmental parameters (TP, TN, DOC, temperature, pH), using data from 460 Norwegian lakes.
2.  Ochromonas was able to feed on all four cyanobacterial strains tested, and grew quickly on all of them. The chrysophyte caused net growth reductions in all three Microcystis strains (the very toxic single-celled strain PCC 7806; the less toxic colony-forming Bear AC and the less toxic single-celled Spring CJ). The effect of Ochromonas was strongest on the Spring CJ strain. Although the effect of Ochromonas grazing on the growth of Pseudanabaena was relatively smaller, it also reduced the net growth of this cyanobacterium significantly.
3. After 4 days of incubation with Ochromonas the total amount of cyanotoxins in the three Microcystis strains was reduced by 91.1–98.7% compared with the controls.
4.  Ochromonas occurred in similar densities across all 460 Norwegian lakes. Microcystis occurred only at higher TN, TP, temperature and pH values, although its density was often several orders of magnitude higher than that of Ochromonas . Ochromonas co-occurred in 94% of the samples in which Microcystis was present.
5. From our study it is not clear whether Ochromonas could control Microcystis blooms in natural lakes. However, our study does demonstrate that Ochromonas usually occurs in lakes with Microcystis , and our small scale experiments show that Ochromonas can strongly reduce the biomass of Microcystis and its toxin content.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

Natural community bioassays to determine the abiotic factors that control phytoplankton growth and succession

SUMMARY. 1. The successional patterns of the dominant phytoplankton species in Lake Maarsseveen (The Netherlands) were very similar in 1980 and 1981. In December/January the diatoms Stephanodiscus hantzschii Grun., Stephanodiscus astraea (Ehr.) Grun. and Asterionella formosa Hass. dominated the algal community (A. formosa had several further population increases during the year). Fragilaria crotonensis Kitt. came to dominance in March/April, followed by the chrysophyte Dinobryon divergens Imhof and the diatom Cyclotella comta (Ehr.) Kütz in May/June. A second appearance of D. divergens was observed in July/August, followed in 1980 by F. crotonensis and a third small increase of D. divergens. In both years S. astraea and S. hantzschii started to grow again in November/December. Cryptophyceae and Chlorophyta were present throughout the year, but did not show a distinct succession. 2. Natural community bioassays, performed under natural light and temperature conditions in a newly developed outdoor bioassay apparatus, showed that phosphate was the major nutrient limiting the growth rate of the phytoplankton. From January till June, during the decline in phosphorus concentration, the diatoms became successively phosphate limited in the sequence: S. hantzschii, S. astraea, F. crotonensis, A. formosa and C. comta. Light limitation was probably the major cause of the relatively late start of F. crotonensis in early spring. 3. D. divergens, increasing after the diatoms from June till September, was stimulated by the addition of a chelator (EDTA). The chelator might stimulate the formation of trace metal species favouring their uptake (e.g. iron). 4. The patterns of succession of the diatoms observed from January till June and from July till December were to a large extent symmetrical. The controlling factors followed opposite trends: declining phosphorus concentrations with increasing irradiance from winter till spring and increasing phosphorus concentrations with decreasing irradiance from summer till late winter.