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In many lakes, the most conspicuous seasonal events are the phytoplankton spring bloom and the subsequent clear-water phase, a period of low-phytoplankton biomass that is frequently caused by mesozooplankton (Daphnia) grazing. In Central European lakes, the timing of the clear-water phase is linked to large-scale climatic forcing, with warmer winters being followed by an earlier onset of the clear-water phase. Mild winters may favour an early build-up of Daphnia populations, both directly through increased surface temperatures and indirectly by reducing light limitation and enhancing algal production, all being a consequence of earlier thermal stratification. We conducted a field experiment to disentangle the separate impacts of stratification depth (affecting light supply) and temperature on the magnitude and timing of successional events in the plankton. We followed the dynamics of the phytoplankton spring bloom, the clear-water phase and the spring peak in Daphnia abundance in response to our experimental manipulations. Deeper mixing delayed the timing of all spring seasonal events and reduced the magnitudes of the phytoplankton bloom and the subsequent Daphnia peak. Colder temperatures retarded the timing of the clear-water phase and the subsequent Daphnia peak, whereas the timing of the phytoplankton peak was unrelated to temperature. Most effects of mixing depth (light) and temperature manipulations were independent, effects of mixing depth being more prevalent than effects of temperature. Because mixing depth governs both the light climate and the temperature regime in the mixed surface layer, we propose that climate-driven changes in the timing and depth of water column stratification may have far-reaching consequences for plankton dynamics and should receive increased attention.
Transient dynamics of pelagic producer-grazer systems in a gradient of nutrients and mixing depths
(2008)
Phytoplankton–grazer dynamics are often characterized by long transients relative to the length of the growing season. Using a phytoplankton–grazer model parameterized for Daphnia pulex with either flexible or fixed algal carbon : nutrient stoichiometry, we explored how nutrient and light supply (the latter by varying depth of the mixed water column) affect the transient dynamics of the system starting from low densities. The system goes through an initial oscillation across nearly the entire light–nutrient supply space. With flexible (but not with fixed) algal stoichiometry, duration of the initial algal peak, timing and duration of the subsequent grazer peak, and timing of the algal minimum are consistently accelerated by nutrient enrichment but decelerated by light enrichment (decreasing mixing depth) over the range of intermediate to shallow mixing depths. These contrasting effects of nutrient vs. light enrichment are consequences of their opposing influences on food quality (algal nutrient content): algal productivity and food quality are positively related along a nutrient gradient but inversely related along a light gradient. Light enrichment therefore slows down grazer growth relative to algal growth, decelerating oscillatory dynamics; nutrient enrichment has opposite effects. We manipulated nutrient supply and mixing depth in a field enclosure experiment. The experimental results were qualitatively much more consistent with the flexible than with the fixed stoichiometry model. Nutrient enrichment increased Daphnia peak biomass, decreased algal minimum biomass, decreased the seston C:P ratio, and accelerated transient oscillatory dynamics. Light enrichment (decreasing mixing depth) produced the opposite patterns, except that Daphnia peak biomass increased monotonously with light enrichment, too. Thus, while the model predicts the possibility of the “paradox of energy enrichment” (a decrease in grazer biomass with light enrichment) at high light and low nutrient supply, this phenomenon did not occur in our experiment.
We independently manipulated mixing intensity (strong artificial mixing vs. background turbulence) and water‐column depth (2 m, 4 m, 8 m, and 12 m) in order to explore their separate and combined effects in a field enclosure experiment. To accentuate the vertical light gradient, enclosures had black walls, resulting in a euphotic depth of only 3.7 m. All enclosures were placed in a well‐mixed water bath to equalize temperature across treatments. Phytoplankton responded to an initial phosphorus pulse with a transient increase in biomass, which was highest in the shallowest, least light‐limited water columns where dissolved mineral phosphorus subsequently became strongly limiting. As a consequence, the depth‐averaged mineral phosphorus concentration increased and the seston carbon (C) : phosphorous (P) ratio decreased with increasing water‐column depth. Low turbulence enclosures became quickly dominated by motile taxa (flagellates) in the upper water column, whereas mixed enclosures became gradually dominated by pennate diatoms, which resulted in higher average sedimentation rates in the mixed enclosures over the 35‐d experimental period. Low turbulence enclosures showed pronounced vertical structure in water columns >4 m, where diversity was higher than in mixed enclosures, suggesting vertical niche partitioning. This interpretation is supported by a primary production assay, where phytoplankton originating from different water depths in low‐turbulence treatments had the relatively highest primary productivity when incubated at their respective depths of origin.
Physical determinants of phytoplankton production, algal stoichiometry, and vertical nutrient fluxes
(2010)
Most phytoplankters face opposing vertical gradients in light versus nutrient supplies but have limited capacities for vertical habitat choice. We therefore explored a dynamical model of negatively buoyant algae inhabiting a one-dimensional water column to ask how water column depth and turbulence constrain total (areal) phytoplankton biomass. We show that the population persistence boundaries in water column depth-turbulence space are set by sinking losses and light limitation but that nutrients are most limiting to total biomass in water columns that are neither too shallow or too weakly mixed (where sinking losses prevail) nor too deep and turbulent (where light limitation prevails). In shallow waters, the most strongly limiting process is nutrient influx to the bottom of the water column (e.g., from sediments). In deep waters, the most strongly limiting process is turbulent upward transport of nutrients to the photic zone. Consequently, the highest total biomasses are attained in turbulent waters at intermediate water column depths and in deep waters at intermediate turbulences. These patterns are insensitive to the assumption of fixed versus flexible algal carbon-to-nutrient stoichiometry, and they arise irrespective of whether the water column is a surface layer above a deep water compartment or has direct contact with sediments.
The Bode catchment (Germany) shows strong land use gradients from forested parts of the National Park (23 % of total land cover) to agricultural (70 %) and urbanised areas (7 %). It is part of the Terrestrial Environmental Observatories of the German Helmholtz association. We performed a biogeochemical analysis of the entire river network. Surface water was sampled at 21 headwaters and at ten downstream sites, before (in early spring) and during the growing season (in late summer). Many parameters showed lower concentrations in headwaters than in downstream reaches, among them nutrients (ammonium, nitrate and phosphorus), dissolved copper and seston dry mass. Nitrate and phosphorus concentrations were positively related to the proportion of agricultural area within the catchment. Punctual anthropogenic loads affected some parameters such as chloride and arsenic. Chlorophyll a concentration and total phosphorus in surface waters were positively related. The concentration of dissolved organic carbon (DOC) was higher in summer than in spring, whereas the molecular size of DOC was lower in summer. The specific UV absorption at 254 nm, indicating the content of humic substances, was higher in headwaters than in downstream reaches and was positively related to the proportion of forest within the catchment. CO2 oversaturation of the water was higher downstream compared with headwaters and was higher in summer than in spring. It was correlated negatively with oxygen saturation and positively with DOC concentration but negatively with DOC quality (molecular size and humic content). A principle component analysis clearly separated the effects of site (44 %) and season (15 %), demonstrating the strong effect of land use on biogeochemical parameters.
Forschungsbericht 2012
(2013)
In shallow aquatic systems, benthic and pelagic primary producers typically compete for light and nutrients along opposing vertical supply axes: pelagic algae shade the benthic habitat; conversely, benthic algae intercept the nutrient flux from the sediment to the pelagic habitat. We present a general framework for analyzing such spatially asymmetric resource competition across habitat boundaries using a mechanistic, dynamical model. We visualize the mechanisms determining the outcome of these cross‐habitat interactions using zero‐net‐growth isoclines, resource supply points, and resource consumption vectors. In extensive invasion analyses, we characterize the abiotic and competitive persistence boundaries of pelagic and benthic primary producers, which are set by environmental factors determining nutrient and light supply and are modified by resource use by the competitor in the respective other habitat.
We note several qualitative differences between cross‐habitat and “classical” within‐habitat resource competition. First, coexistence of cross‐habitat competitors is facilitated by, but does not require niche differentiation with respect to, the utilization of resources. Because each species has a competitive edge for the resource that is supplied from “its” side of the system, a competitor that is inferior in utilizing both resources can sometimes coexist with, or even exclude, a superior competitor. Second, increasing the external supply of one resource (the nutrient) may initially favor both competitors, until a breakpoint is reached where the benthic producer goes abruptly extinct. Finally, whether a given pair of cross‐habitat competitors coexist or shows alternative states may depend on the environment. Specifically, benthic and pelagic algae may coexist at low nutrient and light supply but produce alternative states at high nutrient and light supply. Alternative states are, in turn, promoted by any algal trait combination that increases the spatial asymmetry in resource consumption, i.e., leads to a higher nutrient consumption in the benthic habitat and/or a higher light consumption in the pelagic habitat.
In a first empirical application, we show that predictions from our model give a good fit to published data on benthic and pelagic primary production in temperate and arctic lakes spanning a broad range of nutrient environments.
1. Humic lakes with a high external supply of DOC and low input of nutrients can often support a high biomass of metazoan zooplankton. In such lakes, autotrophic algae compete with bacteria for inorganic nutrients, but bacteria support mixotrophic growth. Consequently, planktonic communities are often dominated by mixotrophic flagellates, while obligate autotrophic phytoplankton occurs in low numbers for extended periods.
2. To test the importance of autotrophic phytoplankton and mixotrophic flagellates as food resources for metazoan grazers and, in turn, the feedback effects of grazers on basal food‐web interactions, we conducted a long‐term experiment where we simulated abiotic resource relationships of humic lakes (high DOC [glucose] and low P input). We examined the population dynamics of Daphnia galeata when inoculated in systems with autotrophic algae only, mixotrophic algae only and a mixture of autotrophic and mixotrophic algae, and how the systems changed after the inoculation of Daphnia. All combinations were run at high‐ and low‐light conditions to analyse the effects of light on food quantity and quality.
3. Daphnia grew to high densities only when mixotrophs were present at high‐light conditions and showed no or only weak growth at low‐light conditions or with autotrophs as the only food source.
4. Autotrophic algae and bacteria showed a strong competition for nutrients. Autotrophic algae were released from competition for nutrients after Daphnia grazed on bacteria, which led to a probable change of the bacteria community to less edible but less competitive taxa. As a consequence, there was a mutualistic interaction between autotrophs and mixotrophs before Daphnia were introduced which turned into competition after Daphnia inoculation.
5. We suggest that mixotrophic flagellates can be a critical resource for cladocerans and thereby also have a cascading effect on higher trophic levels, and cladocerans, in turn, have important indirect effects on basal planktonic food webs; hence, both might affect whole lake ecosystems.
Forschungsbericht 2013
(2014)
Forschungsbericht 2014
(2015)