Tobias Vrede

Fundamental connections among organism c:n:p stoichiometry, macromolecular composition and growth.

Whereas it is acknowledged that the C:N:P stoichiometry of consumers and their resources affects both the structure and the function of food webs, and eventually influences broad-scale processes su ...

Fatty Acid Ratios in Freshwater Fish, Zooplankton and Zoobenthos – Are There Specific Optima?

Two groups of polyunsaturated fatty acids (PUFA), termed omega-3 and omega-6 in food (or here as n-3 and n-6 PUFA, respectively), are essential for all vertebrates and probably also for nearly all invertebrates. The absolute concentrations of the different PUFA are important, as is an appropriate balance between the two. The optimal ratio of n-3/n-6 is not known for most organisms but is anticipated to be more or less species-specific (Sargent et al. 1995). The three most important PUFA in vertebrates are eicosapentaenoic acid (EPA, 20:5n-3), docosahexaenoic acid (DHA, 22:6n-3) and arachidonic acid (ARA, 20:4n-6). Both EPA and ARA are precursors for biologically active eicosanoids that are vital components of cell membranes and play many dynamic roles in mediating and controlling a wide array of cellular activities (Crawford et al. 1989; Harrison 1990; Henderson et al. 1996; see Chap. 9). Since n-3 and n-6 PUFA cannot be synthesized de novo by most metazoans, they must be included in the diet, either as EPA, DHA and ARA, or as their precursors, such as α-linolenic acid (ALA, 18:3n-3, precursor of EPA and DHA) and linoleic acid (LIN, 18:2n-6, precursor of ARA) (Bell et al. 1986; Sargent et al. 1995). Both ALA and LIN are produced in the thylacoid membranes of algae and plants with chlorophyll (Sargent at al. 1987).

Iron Constraints on Planktonic Primary Production in Oligotrophic Lakes

Phototrophic primary production is a fundamental ecosystem process, and it is ultimately constrained by access to limiting nutrients. Whereas most research on nutrient limitation of lacustrine phytoplankton has focused on phosphorus (P) and nitrogen (N) limitation, there is growing evidence that iron (Fe) limitation may be more common than previously acknowledged. Here we show that P was the nutrient that stimulated phytoplankton primary production most strongly in seven out of nine bioassay experiments with natural lake water from oligotrophic clearwater lakes. However, Fe put constraints on phytoplankton production in eight lakes. In one of these lakes, Fe was the nutrient that stimulated primary production most, and concurrent P and Fe limitation was observed in seven lakes. The effect of Fe addition increased with decreasing lake water concentrations of total phosphorus and dissolved organic matter. Possible mechanisms are low import rates and low bioavailability of Fe in the absence of organic chelators. The experimental results were used to predict the relative strength of Fe, N, and P limitation in 659 oligotrophic clearwater lakes (with total phosphorus ≤ 0.2 μM P and total organic carbon < 6 mg C l−1) from a national lake survey. Fe was predicted to have a positive effect in 88% of these lakes, and to be the nutrient with the strongest effect in 30% of the lakes. In conclusion, Fe, along with P and N, is an important factor constraining primary production in oligotrophic clearwater lakes, which is a common lake-type throughout the northern biomes.

Separating physical and physiological effects of temperature on zooplankton feeding rate

Abstract.The effect of temperature and viscosity on the ingestion rate of Daphnia galeata and Chydorus sphaericus was investigated in short-term feeding experiments with 3H-labelled Scenedesmus quadricauda. Viscosity was manipulated independently from temperature by addition of inert polymers in two experiments. In the first experiment, the ingestion rate of D. galeata was found to be 50% lower at 10°C than at 25°C due to the combined effects of increased viscosity and lower temperature. Viscosity alone accounted for 61% of the decline in ingestion rate. In the second experiment, the ingestion rate of a different clone of D. galeata was 46–53% lower at 10°C than at 20°C and changes in viscosity alone accounted for most (>92%) of that decline. The ingestion rate of C. sphaericus was 80% lower at 10°C than at 20°C, and viscosity accounted for 65% of that decline. Thus, viscosity plays an important role in determining the ingestion rate of crustacean zooplankton. Adaptations to life in waters that undergo considerable variation in temperature and viscosity may therefore influence energy budgets and may eventually be of importance for zooplankton species succession, distribution and evolution.

Autochthonous resources are the main driver of consumer production in dystrophic boreal lakes

Dystrophic lakes are widespread in temperate regions and intimately interact with surrounding terrestrial ecosystems in energy and nutrient dynamics, yet the relative importance of autochthonous and allochthonous resources to consumer production in dystrophic lakes remains controversial. We argue that allochthonous organic matter quantitatively dominates over photosynthetic autotrophs in dystrophic lakes, but that autotrophs are higher in diet quality and more important for consumers as they contain essential polyunsaturated fatty acids (PUFA). In a field study, we tested the hypotheses that (1) autochthonous primary production is the main driver for consumer production, despite being limited by light availability and low nutrient supplies, and greater supply of allochthonous carbon, (2) the relative contribution of autotrophs to consumers is directly related to their tissue PUFA concentrations, and (3) methane-oxidizing bacteria (MOB) provide an energy alternative for consumers. Pelagic and benthic consumer taxa representing different trophic levels were sampled from five dystrophic lakes: isopod Asellus aquaticus, megalopteran Sialis lutaria, dipteran Chaoborus flavicans, and perch Perca fluviatilis. Based on carbon and nitrogen stable isotopes, the relative contributions of autochthonous (biofilms and seston) and allochthonous (coarse particulate and dissolved organic matter) resources and MOB to these taxa were 47-79%, 9-44% and 7-12% respectively. Results from fatty acid (FA) analyses show that the relative omega3-FA and PUFA concentrations increased with trophic level (Asellus < Sialis and Chaoborus < Perca). Also, eicosapentaenoic-acid (EPA), omega3-FA and PUFA concentrations increased with the autochthonous contribution in consumers, i.e., a 47-79% biofilm and/or seston diet resulted in tissue EPA of 4.2-18.4, omega3 FAs of 11.6-37.0 and PUFA of 21.6-61.0 mg/g dry mass. The results indicate that consumers in dystrophic lakes predominantly rely on energy from autotrophs and that their PUFA concentrations are dependent on the relative contribution of these autochthonous resources. The limited energy support from MOB suggests they are not negligible and are potentially an integral part of the food webs. Our findings show that autochthonous resources are the main driver of secondary production even in dystrophic lakes and offer new insights into the functioning of these ecosystems.

Transfer of bacterial production based on labile carbon to higher trophic levels in an oligotrophic pelagic system

Additions of labile organic carbon (C) enhanced bacterial production (BP) and were associated with increases in crustacean zooplankton and planktivorous fish biomasses. This was shown in a mesocosm ...

Zooplankton Feeding on the Nuisance Flagellate Gonyostomum semen

The large bloom-forming flagellate Gonyostomum semen has been hypothesized to be inedible to naturally occurring zooplankton due to its large cell size and ejection of long slimy threads (trichocysts) induced by physical stimulation. In a grazing experiment using radiolabelled algae and zooplankton collected from lakes with recurring blooms of G. semen and lakes that rarely experience blooms, we found that Eudiaptomus gracilis and Holopedium gibberum fed on G. semen at high rates, whereas Daphnia cristata and Ceriodaphnia spp. did not. Grazing rates of E. gracilis were similar between bloom-lakes and lakes with low biomass of G. semen, indicating that the ability to feed on G. semen was not a result of local adaptation. The high grazing rates of two of the taxa in our experiment imply that some of the nutrients and energy taken up by G. semen can be transferred directly to higher trophic levels, although the predominance of small cladocerans during blooms may limit the importance of G. semen as a food resource. Based on grazing rates and previous observations on abundances of E. gracilis and H. gibberum, we conclude that there is a potential for grazer control of G. semen and discuss why blooms of G. semen still occur.

Community structure in boreal lakes with recurring blooms of the nuisance flagellate Gonyostomum semen

Blooms of the nontoxic raphidophyte Gonyostomum semen have shown a recent increase in frequency and distribution in the Fennoscandian region. Due to large cell size and several grazer-avoidance strategies, G. semen is hypothesized to be inedible for most zooplankton species and therefore may constitute a bottleneck for the transfer of energy and nutrients in pelagic food webs. Repression of other phytoplankton through increased competition and induced mortality could further exacerbate this effect. In a field study of four lakes with recurring blooms of G. semen and four lakes without blooms, we found significant differences in community structure between the two lake groups during the bloom period. Bloom-lakes had lower biovolumes of small chrysophytes and chlorophytes and zooplankton assemblages were predominated by small, potentially bacterivorous cladocerans, suggesting a limited availability of edible phytoplankton and an increased importance of microbial pathways during G. semen blooms. Low biovolumes of large cladocerans in bloom-lakes may be due to interference of G. semen with filter feeding. Moreover, high abundances of the phantom midge Chaoborus flavicans in bloom-lakes suggest that the flow of energy and nutrients is directed more towards this invertebrate predator than fish. This could have negative impacts on fish populations, especially if bloom periods are prolonged.

Contrasting “top-down” effects of crustacean zooplankton grazing on bacteria and phytoflagellates

The combined effects of grazing and nutrient regeneration by Daphnia and Eudiaptomus on the growth of Rhodomonas and heterotrophic bacteria was assessed experimentally. The responses of Rhodomonas and bacteria to the grazers were measured as net specific growth rate over the entire experimental periods, as well as production and specific production at the end of the experiments. Both zooplankton species had a negative effect on Rhodomonas net specific growth rate due to grazing and a positive effect on specific primary production due to nutrient regeneration. Daphnia had no effect on bacterial net specific growth rate, bacterial production or specific bacterial production in one of two experiments. In the other experiment, however, both bacterial growth rate and production decreased as a result of grazing. Furthermore, Daphnia had a negative effect on specific bacterial production, but Eudiaptomus had a positive effect on all bacterial parameters due to nutrient regeneration, probably of phosphorus. Positive effects of copepods on bacterial growth has previously been attributed to trophic cascades via protozoa. However, the present experiments show that regeneration of nutrients, especially phosphorus, may account for a large part of the stimulation of bacterial growth.

Responses in zooplankton populations to food quality and quantity changes after whole lake nutrient enrichment of an oligotrophic sub-alpine reservoir

Abstract.To sustain production of higher trophic levels in oligotrophic systems it is important that the trophic transfer of energy and nutrients is efficient. The phytoplankton-zooplankton interface is of specific interest since nutritional constraints can decouple energy flow in this step. Increased nutrient loading to oligotrophic systems with initially low abundance of high quality phytoplankton could induce changes in seston composition that reduces the nutritional value for zooplankton. We carried out a whole lake enrichment experiment for five years in two ultraoligotrophic subalpine hydroelectric power reservoirs in Sweden. The first year was an untreated reference year. Phosphorus and nitrogen were added to Lake Stora Mjölkvattnet during the following four years, and upstream Lake Burvattnet was used as an untreated reference lake. The phosphorus content of seston in the experimental lake increased in the years of fertilization and seston phosphorus to carbon ratios (atomic) were non-limiting for zooplankton growth. Decreasing concentrations of phosphorus in the reference lake lead to low phosphorus to carbon ratios that probably affected zooplankton growth negatively. The seston fatty acid concentrations and phytoplankton composition indicated good food quality in both lakes. The phytoplankton increased in the experimental lake despite an increase in zooplankton biomass. Some changes in the relative contributions of plankton species occurred but the same species were present. The crustacean zooplankton community composition shifted towards smaller species during the latter years in the experimental lake, indicating increased predation pressure from fish. A major result of this whole-lake ecosystem fertilization experiment is that gentle fertilization can significantly boost phytoplankton production while food quality remains high, and plankton community composition is not substantially altered.