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The productivity and trophic structure of aquatic ecosystems is the result of an interplay between bottom-up and top-down forces that operate both within and across the benthic and pelagic compartments of lake food webs. Contemporary and projected climate changes urge the question how this interplay will be affected by increasing inputs of terrestrial derived, dissolved organic matter (‘browning’) and warming. We addressed this issue by exploring how browning and warming affect the behavior of a relatively simple, conceptual model of a shallow lake food web that is compartmentalized into, dynamically coupled, benthic and pelagic components (abiotic resources, primary producers, grazers, and carnivores). We compared model expectations with the results of a factorial manipulation of browning and warming in a replicated, large-scale field experiment. Both the model and the experiment suggest that browning affects the food web from the bottom-up by reducing light supply to the benthic habitat and increasing nutrient supply to the pelagic habitat, with concomitant decreases of benthic and increases of pelagic primary and secondary production. The model also predicts that warming effects should primarily operate via relaxed top-down control by top consumers in the more productive of the two habitats. The latter was only partially supported by the experimental data, possibly because the model still lacks one or two important trophic links, such as the one from pelagic producers to benthic deposit feeders. We propose that our coupled benthic-pelagic food web model provides a useful conceptual starting point for future theoretical and empirical studies of the impacts of environmental changes on shallow lakes.

Biological stoichiometry is an approach that focuses on the balance of elements in biological interactions. It is a theory that has the potential to causally link material processes at all biological levels—from molecules to the biosphere. But the lack of a coherent operational framework has so far restricted progress in this direction. Here, we provide a framework to help infer how a stoichiometric imbalance observed at one level impacts all other biological levels. Our framework enables us to highlight the areas of the theory in need of completion, development and integration at all biological levels. Our hope is that this framework will contribute to the building of a more predictive theory of elemental transfers within the biosphere, and thus, to a better understanding of human-induced perturbations to the global biogeochemical cycles.

The majority of lakes on Earth are shallow, unproductive and located at high latitudes. These lakes are experiencing big changes due to climate change, where two environmental drivers operate simultaneously, browning and warming. How they affect lake ecosystems is not well understood. Here, I addressed this issue by using a theoretical and an experimental approach. In particular, I generated model predictions and compared them with the results of a realistic large-scale experiment, where browning and warming were manipulated in a factorial design. In addition, model outcomes were compared with data from 12 unproductive lakes sampled along a gradient of browning. Another novelty of my thesis is that it integrates benthic and pelagic food web components in the model and experimental approaches. I found that browning affected the resources availability for benthic and pelagic producers in the model and in the experiment. With browning, benthic primary producers became increasingly light limited and declined, while pelagic producers became less nutrient limited and increased. Pelagic nutrient limitation was alleviated by two non-exclusive mechanisms. Browning directly enriched the water with nutrients, and browning indirectly increased the nutrient flowing from the sediment to the pelagic habitat via suppression of benthic producers. To tease apart these two mechanisms I applied structural equation modeling (SEM). The indirect evidence by SEM suggests that both mechanisms contributed equally to the pelagic nutrient concentration in the experiment. Interestingly, a model food web with only primary producers shows similar qualitative behavior as a food web with grazers and carnivores included. This happens because carnivorous fish exert strong top-down control in the more productive habitat, which relaxes grazing pressure on primary producers and increases resource limitation in the adjacent habitat. Biomass of benthic and pelagic consumers followed the same pattern as their resources. The lake data were largely congruent with model expectations and supported the findings of the experiment. Furthermore, the model also predicted a negative relationship between total phosphorus and both primary and fish production, which was observed across the 12 lakes. Warming effects were more complex. The model predicts that warming effects should depend on browning and are expected to be strongest in the more productive of the two (benthic and pelagic) habitats. For example, at low levels of browning the biomasses of benthic algae and fish are expected to decline with warming, which was observed in the experiment. In contrast, observed warming effects at high levels of browning deviated from model expectations. The mechanisms by which browning and warming interactively affect lake food webs are still poorly understood. This thesis offers a conceptual foundation for their further study through the integration of within- and between-habitat interactions.

In shallow lakes, pelagic and benthic producers engage in spatially asymmetrical resource competition. Pelagic producers intercept the flux of light to the benthic habitat and benthic producers intercept the flux of sediment-derived nutrients to the pelagic habitat. In boreal and subarctic regions, climate change is affecting this interaction both directly through warming and indirectly through increased loading with colored dissolved organic matter (cDOM) from the catchment (brownification). We use a dynamical ecosystem model to explore the consequences of these changing environmental conditions for lake primary production and compare model predictions with the results of an experiment in which we manipulated water temperature and cDOM supply in a 2x2 factorial design. The experiment was performed in field mesocosms large enough to harbor reproducing fish populations and was run over an entire growing season. In agreement with model predictions, benthic algal production and biomass declined and pelagic algal production and biomass increased with browning. Pelagic nutrient concentrations diverged over time between low and high cDOM treatments, suggesting that browning alleviated pelagic algal nutrient limitation by shading benthic competitors and preventing them from intercepting the release of nutrients from the sediment. Warming considerably reduced benthic and pelagic algal production as well as pelagic algalbiomass and total phosphorus. The warming results are only in partial accordance with model expectations, but can be explained by an indirectly inferred, positive response of macrophyte production (which was not included in the model) to warming. Our study suggests that lake ecosystem responses to climate change are mediated by cross-habitat feedbacks between benthic and pelagic producers.

Global warming is expected to influence lake gross primary production (GPP) and ecosystem respiration (R) by increasing water temperature and terrestrial export of organic material and inorganic nutrients from the catchment. We experimentally tested the effects of warming (3 A degrees C) and natural humic river runoff, separately and in combination, on habitat-specific and whole ecosystem net ecosystem production (NEP = GPP - R) in replicated large scale (136 m(3)) experimental pond ecosystems over one open water season. Pelagic NEP was reduced by warming and increased with humic river water addition. Littoral NEP (benthos, macrophytes, periphyton) showed an opposite pattern with increasing NEP following warming and decreasing NEP following humic river water addition. These changes were a result of changes in GPP with warming (negative in pelagic, positive in littoral) and with humic water addition (positive in pelagic, negative in littoral), while no effects were observed on pelagic respiration. As a result of the counteracting effects on NEP in pelagic and littoral habitats, whole ecosystem NEP was not affected by the treatments. The study suggests that climate mediated changes in temperature and river runoff have relatively small effects on the overall metabolic balance of shallow aquatic ecosystems but there may be large habitat-specific effects.

Climate change is expected to not only raise water temperatures, but also to cause brownification of aquatic ecosystems via increased inputs of terrestrial dissolved organic matter. While efforts have been made to understand how increased temperature and brownification separately influence aquatic food webs, their interactive effects have been less investigated. Further, although climate change effects on aquatic ecosystems likely will propagate to terrestrial consumers via changes in aquatic insect emergence, this has rarely been studied. We investigated the effect of climate change on aquatic insect emergence, in a large-scale outdoor pond facility where 16 sections - each containing natural food webs including a fish top-consumer population - were subjected to warming (3 degrees C above ambient temperatures) and/or brownification (by adding naturally humic stream water). Aquatic insect emergence was measured biweekly over 18weeks. We found no effect of warming or brownification on total emergent insect dry mass. However, warming significantly reduced the number of emergent Chironomidae, while numbers of larger taxa, Trichoptera and Ephemeroptera, remained unchanged. On average, 57% and 58% fewer Chironomidae emerged from the warmed clear and humic pond sections, respectively. This substantial decrease in emergent Chironomidae resulted in a changed community structure and on average larger individuals emerging from warm sections as well as from humic sections under ambient conditions. There was also a weak influence of fish biomass on the size structure of emergent aquatic insects, with a positive relationship between individual insect size and total fish biomass, but effects of fish were clearly subordinate to those of warming. Climate change impacts on aquatic systems can have widespread consequences also for terrestrial systems, as aquatic insects are ubiquitous and their emergence represents an important resource flow from aquatic to terrestrial environments. While we found that neither warming nor brownification quantitatively changed total aquatic insect emergence biomass, the warming-induced decrease in number of emergent Chironomidae and the subsequent increase in average body size will likely impact terrestrial consumers relying on emergent aquatic insect as prey.

Flows of energy and matter across habitat boundaries can be major determinants of the functioning of recipient ecosystems. It is currently debated whether terrestrial dissolved organic matter (tDOM) is a resource subsidy or a resource subtraction in recipient lakes. In a long-term field experiment, pelagic phosphorus concentration and aquatic primary production increased with increasing tDOM input, suggesting that tDOM acted primarily as a direct nutrient subsidy. Piecewise structural equation modeling supports, however, an approximately equally important role for a second mechanism: colored tDOM acted also as a resource subtraction by shading benthic algae, preventing them from intercepting nutrients released across the sediment-water interface. Inhibition of benthic algae by colored tDOM thus indirectly promoted pelagic algae and whole-ecosystem primary production. We conclude that cross-ecosystem terrestrial DOM inputs can modify light and nutrient flows between aquatic habitats and alter the outcome of resource competition between benthic and pelagic producers. These results are particularly relevant for shallow northern lakes, which are facing increased tDOM runoff with climate change.