Global change is altering the community composition, variability, and behavior of organisms in a diverse suite of ecosystems. Because of climate change and eutrophication, freshwater lakes and reservoirs are experiencing an increase in low dissolved oxygen concentrations (hypoxia) in their bottom waters (hypolimnion), which can disrupt ecological communities. Zooplankton, important aquatic organisms for regulating water quality and food webs, are one group of organisms affected by hypoxia since zooplankton need oxygen to respire. My research shows that hypoxia may disrupt zooplankton behavior and increase the variability of zooplankton communities. Zooplankton ubiquitously exhibit diel vertical migration, where the majority of the population resides in the hypolimnion during the daytime to escape predation from fish and damage from ultraviolet radiation. At night, many zooplankton ascend to the surface waters to feed on phytoplankton, when there is decreased risk of predation and radiation. My results from intensive 24-hour sampling campaigns suggest that hypolimnetic hypoxia may alter zooplankton migration, biomass, and behavior, which may in turn exacerbate water quality degradation due to the critical role zooplankton play in freshwater ecosystems. In addition, field surveys in four reservoirs over three years revealed that hypoxia may increase the variability of zooplankton communities compared to oxic conditions. Consequently, as lakes and reservoirs experience increased extent and duration of hypoxia in the future, it is critical to understand how more variable zooplankton communities alter freshwater ecosystem functioning. / Ph. D. / The globalization and extensive use of habitats by humans has created stress on other organisms in many ecosystems; the interactions, migration patterns, and composition of species have been altered globally because of humans. In freshwater lakes and reservoirs, warmer water temperatures and increased nutrient pollution have resulted in decreased oxygen concentrations in many waterbodies. Decreased oxygen concentrations can affect the organisms living in these environments that need oxygen to survive, which can disrupt the water quality and function of the ecosystem. Zooplankton, or animal plankton or animal drifters, are one group of organisms affected by low oxygen conditions. Zooplankton play an important role in aquatic ecosystems. Zooplankton eat phytoplankton (at the base of the food chain) and are consumed by fish; therefore, any changes to zooplankton populations from low oxygen concentrations can affect water quality and fisheries. My research shows that low oxygen concentrations affect the migration patterns of zooplankton. Zooplankton generally migrate vertically in the water column of a lake and reservoir over each 24-hour period to optimize their proximity to phytoplankton (their food source), while minimizing their chance of being eaten by fish. I found that low oxygen concentrations disrupt this cycle; zooplankton largely do not vertically migrate under low oxygen concentrations. My results from intensive 24-hour sampling campaigns and sampling over multiple summer periods suggest that low oxygen concentrations disrupt zooplankton migration and behavior, which may exacerbate water quality issues in lakes and reservoirs.

Climate change affects lake ecosystems in many ways by direct and indirect temperature impacts on hydrophysical structure, hydrology, chemical cycles and on biological interactions. The identification of these climate signals can be complicated and superimposed by other environmental changes, such as land use changes. The aim of this thesis was to seperate temperature effects from effects of management and other environmental influences and to understand the underlying complex processes causing significant changes of ecosystem states. This understanding is important for decisions of lake and reservoir managers to counteract unfavorable consequences of climate change. For the recent study, long-term data of meteorological, hydrophysical and biological variables (phytoplankton, crustacean zooplankton, fish) of the German drinking water reservoir Saidenbach were analysed. Based on this long-term data set, strong indicators for climate induced changes have been identified. In particular, increasing water temperatures since 1975, earlier break up of ice covers and an altered hydrophysical structure could be detected. Thereby, stratification stability increased and turbulent mixing decreased in summer. The water of the reservoir warmed more than the tributaries resulting in a trend to deeper entrainment of the inflows. As further indicators of climate change, an increased annual biomass of phytoplankton and species shifts in the phytoplankton community in spring and summer had been observed. During the spring mass development, the diatom Aulacoseira subarctica became dominant in recent years with warm winters and early ice-out. Its unusual spatial pattern with occurrences in aphotic depths could be explained by easier resuspension compared to other diatoms. By being resuspended first and establishing a high inoculum, A. subarctica profits from an earlier ice-out and earlier full circulation. In spite of a reduced nutrient loading to the epilimnion, in summer, the diatom Fragilaria crotonensis was displaced increasingly by cyanobacteria. This species shift could be explained well by the hydrophysical regime shift. Although, the annual total phytoplankton biomass increased since 1990, the crustacean zooplankton in Saidenbach reservoir did not seem to profit from improved food resources. To the contrary, Daphnia abundances reduced tremendously. We could show that the influcence of fish stocks were underestimated. The stocked silvercarp may have contributed up to 70 % of the total zooplanktivorous fish biomass which had a temperature and density dependent effect. The faster growth of Daphnia at higher temperatures could not compensate for the more actively grazing fish when stock of zooplankitvorous fish was too high. Still, temperature was identified as the most important factor that explained 29 % of the zooplankton phenology, while the second most important predictors were zooplanktivorous fish biomass and nutrient loading, explaining 18 % of the variance. The importance of submerged macrophytes in shallow lakes is well investigated. To increase also the understanding of their impact and their role during climate change on water quality in deep lakes, a model for stratified lakes that includes submerged macrophytes was developed. The simulations showed that macrophyte effects were mainly positive for water quality and macrophytes in deep lakes were able to potentially reduce summer phytoplankton, especially cyanobacteria by 50 % in 11 m deep and still by 15 % in 100 m deep oligotrophic lakes. Nutrient competition with phytoplankton contributed most to this macrophyte effect. In conclusion, for deep lake restoration the re-establishment of submerged macrophytes might be as important as for shallow lakes. The full lake model includes hydrophysical and ecological submodules and thus will allow further comprehensive climate simulations and the evaluation of the effectivity of adaptive strategies and scenarios for deep lakes and reservoirs. / Der Klimawandel beeinflusst Seeökosysteme vielfältig durch direkte und indirekte Temperatureffekte auf die hydrophysikalische Struktur, die Hydrologie, chemische Kreisläufe und biologische Interaktionen. Die Identifikation von Klimasignalen kann durch Landnutzungs- und weitere Umweltveränderungen überlagert werden. Ziel dieser Arbeit war es, Temperatureffekte von Effekten zu unterscheiden, die durch Gewässermanagement oder anderen Umweltveränderungen verursacht werden. Weiterhin sollten komplexe Prozesse verstanden werden, die zu signifikanten Veränderungen in Seeökosystemen führen. Dieses Verständnis ist für Talsperren- und Gewässermanager von besonderer Bedeutung, um ungewünschten Folgen des Klimawandels entgegenwirken zu können. Für die Studie wurden Langzeitdaten für meteorologische, hydrophysikalische und biologische Variablen (Phytoplankton, Crustaceen-Zooplankton, Fischbesatz) der Talsperre Saidenbach ausgewertet. Mehrere Indikatoren für die Auswirkungen des Klimawandels konnten basierend auf diesen Daten identifiziert werden. Diese gehören insbesondere eine Erhöhung der Wassertemperatur seit 1975, zeitigere Eisaufbrüche und eine veränderte Schichtungsstruktur. Dabei wurde im Sommer die Schichtungsstabilität höher und die Intensität der Durchmischung geringer. Das Wasser in der Talsperre erwärmte sich stärker als die Zuflüsse, wodurch diese sich zunehmend tiefer und unterhalb der Thermokline einschichten. Eine zunehmende jährliche Phytoplanktonbiomasse und Artenwechsel innerhalb der Planktongemeinschaft sind weitere Indikatoren für Klimafolgen. Die Kieselalge Aulacoseira subarctica erlangte in Jahren mit warmen Wintern und zeitigen Eisaufbrüchen eine Dominanz während der Frühjahrsentwicklung. Das ungewöhnliche Auftreten dieser Art in aphotischen Tiefen konnte durch eine leichtere Resuspension im Vergleich zu anderen Kieselalgen erklärt werden. Durch zeitige Resuspension gleich nach Eisaufbruch kann A. subarctica ein hohes Inokulum etablieren. Trotz reduzierter Nährstoffzufuhr ins Epilimnion wurde im Sommer die Kieselalge Fragilaria crotonensis vermehrt durch Cyanobakterien verdrängt, was jedoch gut mit den Änderungen der hydrophysikalischen Struktur erklärt werden kann. Obwohl seit 1990 insgesamt mehr Phytoplankton zur Verfügung steht konnte das Crustaceen- Zooplankton in der Talsperre Saidenbach nicht von diesen verbesserten Nahrungsbedingungen profitieren. Die Daphnia-Abundanzen waren seit 1990 stark verringert. Es konnte gezeigt werden, dass der Einfluss des Fischbesatzes unterschätzt wurde und die künstlich ins System eingebrachten Silberkarpfen zeitweise bis zu 70% der zooplanktivoren Fischbiomasse ausgemacht haben. Zoo- planktivore Fische haben einen temperatur- und dichteabhängigen Effekt auf das Zooplankton. Das schnellere Populationswachstum von Daphnia kann bei hohem Besatz mit zooplanktivoren Fischen nicht die erhöhte Fraßaktivität der Fische ausgleichen. Dennoch konnte Temperatur als der wichtigste Einflussfaktor auf die Phänologie des Zooplanktons identifiziert werden, gefolgt von zooplanktivorem Fisch und Nährstoffzufuhr. Die Wichtigkeit von submersen Makrophyten in Flachseen ist gut untersucht. Um den Einfluss von Makropyhten auf die Wasserqualität während des Klimawandels auch in tiefen Seen abschätzen zu können, wurde ein komplexes Seemodell entwickelt. Die Simulationen zeigen, dass sich Makrophyten hauptsächlich positiv auf die Wassergüte auswirken und zur Reduktion von Phytoplankton, insbesondere von Cyanobakterien im Sommer, beitragen. In einem 11 m tiefen See betrug die Reduktion 50 %, in 100 m tiefen oligotrophen Seen immer noch 15 %. Die Konkurrenz um Nährstoffe mit dem Phytoplankton war dabei die ursächliche Makrophyten-Plankton-Interaktion. Submerse Makrophyten könnten für die Restaurierung von tiefen Seen folglich genauso wichtig sein, wie für Flachseen. Das komplette Seemodell enthält hydrophysikalische und ökologische Module und ermöglicht damit weitere umfassende Simulationen zur Untersuchung der Auswirkungen des Klimawandels und zur Evaluation von Adaptionsstrategien für Seen und Talsperren.

Um dos principais problemas relacionados aos estudos limnológicos realizados manualmente em laboratório consiste no grande esforço, tempo de análise e trabalho especializado necessário. Esses fatores limitam a quantidade de amostras possíveis de serem analisadas em um determinado estudo, já que recursos sejam eles reagentes, recursos financeiros ou tempo são limitados. No presente trabalho foi utilizado um sistema semi-automatizado de medidas de variáveis físicas e químicas da água. O sistema é composto por uma sonda multi-parâmetro (Horiba U-22) e um sistema de posicionamento global (GPS) acoplados a um micro-computador, que realizam medidas georeferenciadas em curtos intervalos de tempo, permitindo um rastreamento horizontal das características da água. Foi ainda desenvolvido um sistema semi-automático para coleta fracionada da comunidade planctônica por meio de bomba de sucção operada por bateria e filtro coletor com rede de plâncton de diferentes aberturas de malha. O material coletado foi fotografado por meio de sistema de aquisição digital de imagens (microscópio Zeiss equipado com câmera AxionCan). Neste trabalho foi produzido um software (Planktonscan) que a partir da análise das imagens capturadas permite produzir dados com estimativas das medidas e dimensões dos organismos, calcular biovolumes e, utilizando fatores de conversão, estimar os valores de biomassa. O software apresenta uma interface para identificação, calcula a densidade dos organismos e produz relatório gráfico com informações sobre os organismos individuais e sobre a comunidade. Os equipamentos e o software foram testados em análises limnológicas e amostragem de plâncton no reservatório do Monjolinho, São Carlos, SP, em dezembro de 2005. Os resultados obtidos foram comparados com os disponíveis na literatura e demonstraram a aplicabilidade do sistema. / A major problem associated with the study of planktonic communities lies on the difficulties of analyzing the collected material, a long time-consuming procedure. Biomass determination is also a step requiring great effort and is subjected to large errors. In the present work a semi-automated system for measuring physical and chemical variables in the water was developed. The system is made up by a flow-pump, a multi-parameter probe and a global positioning system coupled to a microcomputer that performs measurements at short time intervals, allowing a horizontal tracking of the water quality, in much shorter times than traditional methods. Another semi-automated device was developed for collecting separate plankton size fractions. It uses a battery operating suction-pump coupled to a filter with different mesh nets. The collected materials are then submitted to image computer acquisition (Axion Vision Zeiss System). Additionally, in this study a software was produced (Planktonscan), that taking the measures of individuals dimensions (length, width and height) calculates biovolume and using conversion factors calculate the biomass for each zooplankton organism identified in the sample. Both systems were tested, regarding the measurement of limnological variables and plankton sampling, in the Monjolinho Reservoir, SP. The performance was good, resulting in a larger number of points sampled (60) in a shorter sampling time (1 hour) than those usually required. The biomass results provided by Planktonscan software were compared to data from literature, obtained by the traditional gravimetric method for dry weight determination and also with data generated from the use of mathematical models (length dry-weight regressions) available. The results were expressed as species population densities, biomasses and size spectra, evidencing the applicability of the models here developed.

Die Gattung Daphnia nimmt eine zentrale Position im pelagischen Nahrungsnetz vieler Standgewässern (Seen, Talsperren) ein. Aufgrund hoher Filtrierleistungen sind Daphnien für das integrierte Gewässermanagement, z.B. im Rahmen einer Nahrungskettenmanipulation (Biomanipulation), von großem Interesse. Ihre relativ einfache Kultivierbarkeit machte sie außerdem zu einen weit verbreiteten Modellorganismus limnologischer Grundlagenforschung. Das Ziel dieser Arbeit besteht darin, ein umfassendes Modellsystem für die Gattung Daphnia zu entwickeln, das wesentliche Kenntnisse der Ökologie von Daphnia miteinander verknüpft, um eine Vernetzung des Wissens zu realisieren. Bestehende Modellansätze konnten diese Aufgabe nur unzureichend erfüllen, weil sie in der Regel einem "Problem-orientierten Ansatz" folgen und somit sehr unterschiedliche inhaltliche Ausrichtungen und verschiedene Modellstrukturen aufweisen. Eine Kopplung bzw. ein direkter Vergleich dieser Modelle ist deshalb in der Regel schwierig. Dieses Modellsystem wurde entwickelt, um ein Instrument zur Synthese des vorhandenen Prozesswissens bereitzustellen, und nicht, um ein spezifisches, abgegrenztes Problem zu bearbeiten; es steht somit diametral zu existierenden Ansätzen. Aus diesem Grunde wird der gewählte Ansatz als ein "Spezies-orientierter Ansatz" bezeichnet. Drei Eigenschaften charakterisieren diesen Ansatz. (i) Das Modellsystem umfasst mehrere hierarchische Ebenen biologischer Organisation, wobei der Schwerpunkt auf den Ebenen des Individuums und der Population liegt. Interaktionen zwischen diesen Organisationsebenen sind ausgeprägt und in Feld- und Laborstudien gut belegt, in existierenden Modellen aber noch unzureichend miteinander verknüpft. (ii) Da eine quantitative Interpretation der Ergebnisse beabsichtigt ist, wurden die Modellausgaben einer umfangreichen Validation an unabhängigen und bereits publizierten Daten unterzogen. (iii) Das Modellsystem zeichnet sich durch einen geschachtelten Aufbau aus (nested design), wodurch eine einfache Erweiterbarkeit des Modells oder auch die separate Anwendung bestimmter Modellkomponenten gewährleistet ist. / The scope of this thesis was to develop a comprehensive model system of the genus Daphnia, a key organism in the pelagic food web of lakes and reservoirs and a widely used model organism in experimental and theoretical ecology. Although its central role in applied and basic research in aquatic ecology is obvious, there are still fundamental problems in modelling the observed dynamics of Daphnia (for details see chapter 2). Therefore, a basic motivation of this work was to use scientific results obtained in independently conducted research for developing a model that brings these results into context. Instead of following a "problem-oriented" paradigm applicable to a single, well defined problem or scientific hypothesis, the underlying concept of the emerging model system was considered to be "species-oriented". Thus, various relevant processes are included into the framework in order to simulate the dynamics of daphnids displayed on different levels of biological organization. To facilitate its application to various problems in ecological research on the genus Daphnia, the model system fulfills the following three important properties: (1) model outputs are thoroughly validated on experimental data in order to guarantee sound quantitative outputs of the model system (2) the system spans over different levels of biological organization with special emphasis laid upon the individual level and the population level (3) the model´s architecture follows a nested design with a defined individual level model that is integrated into a population level model The whole model system is able to describe an individual´s development over time on basis of physiological properties of the organism and, furthermore, how these individual level processes interact with the dynamics on the population level. Due to its nested design, applications of separate submodels (e.g. the individual-level model) are possible.

Climate change affects lake ecosystems in many ways by direct and indirect temperature impacts on hydrophysical structure, hydrology, chemical cycles and on biological interactions. The identification of these climate signals can be complicated and superimposed by other environmental changes, such as land use changes. The aim of this thesis was to seperate temperature effects from effects of management and other environmental influences and to understand the underlying complex processes causing significant changes of ecosystem states. This understanding is important for decisions of lake and reservoir managers to counteract unfavorable consequences of climate change. For the recent study, long-term data of meteorological, hydrophysical and biological variables (phytoplankton, crustacean zooplankton, fish) of the German drinking water reservoir Saidenbach were analysed. Based on this long-term data set, strong indicators for climate induced changes have been identified. In particular, increasing water temperatures since 1975, earlier break up of ice covers and an altered hydrophysical structure could be detected. Thereby, stratification stability increased and turbulent mixing decreased in summer. The water of the reservoir warmed more than the tributaries resulting in a trend to deeper entrainment of the inflows. As further indicators of climate change, an increased annual biomass of phytoplankton and species shifts in the phytoplankton community in spring and summer had been observed. During the spring mass development, the diatom Aulacoseira subarctica became dominant in recent years with warm winters and early ice-out. Its unusual spatial pattern with occurrences in aphotic depths could be explained by easier resuspension compared to other diatoms. By being resuspended first and establishing a high inoculum, A. subarctica profits from an earlier ice-out and earlier full circulation. In spite of a reduced nutrient loading to the epilimnion, in summer, the diatom Fragilaria crotonensis was displaced increasingly by cyanobacteria. This species shift could be explained well by the hydrophysical regime shift. Although, the annual total phytoplankton biomass increased since 1990, the crustacean zooplankton in Saidenbach reservoir did not seem to profit from improved food resources. To the contrary, Daphnia abundances reduced tremendously. We could show that the influcence of fish stocks were underestimated. The stocked silvercarp may have contributed up to 70 % of the total zooplanktivorous fish biomass which had a temperature and density dependent effect. The faster growth of Daphnia at higher temperatures could not compensate for the more actively grazing fish when stock of zooplankitvorous fish was too high. Still, temperature was identified as the most important factor that explained 29 % of the zooplankton phenology, while the second most important predictors were zooplanktivorous fish biomass and nutrient loading, explaining 18 % of the variance. The importance of submerged macrophytes in shallow lakes is well investigated. To increase also the understanding of their impact and their role during climate change on water quality in deep lakes, a model for stratified lakes that includes submerged macrophytes was developed. The simulations showed that macrophyte effects were mainly positive for water quality and macrophytes in deep lakes were able to potentially reduce summer phytoplankton, especially cyanobacteria by 50 % in 11 m deep and still by 15 % in 100 m deep oligotrophic lakes. Nutrient competition with phytoplankton contributed most to this macrophyte effect. In conclusion, for deep lake restoration the re-establishment of submerged macrophytes might be as important as for shallow lakes. The full lake model includes hydrophysical and ecological submodules and thus will allow further comprehensive climate simulations and the evaluation of the effectivity of adaptive strategies and scenarios for deep lakes and reservoirs. / Der Klimawandel beeinflusst Seeökosysteme vielfältig durch direkte und indirekte Temperatureffekte auf die hydrophysikalische Struktur, die Hydrologie, chemische Kreisläufe und biologische Interaktionen. Die Identifikation von Klimasignalen kann durch Landnutzungs- und weitere Umweltveränderungen überlagert werden. Ziel dieser Arbeit war es, Temperatureffekte von Effekten zu unterscheiden, die durch Gewässermanagement oder anderen Umweltveränderungen verursacht werden. Weiterhin sollten komplexe Prozesse verstanden werden, die zu signifikanten Veränderungen in Seeökosystemen führen. Dieses Verständnis ist für Talsperren- und Gewässermanager von besonderer Bedeutung, um ungewünschten Folgen des Klimawandels entgegenwirken zu können. Für die Studie wurden Langzeitdaten für meteorologische, hydrophysikalische und biologische Variablen (Phytoplankton, Crustaceen-Zooplankton, Fischbesatz) der Talsperre Saidenbach ausgewertet. Mehrere Indikatoren für die Auswirkungen des Klimawandels konnten basierend auf diesen Daten identifiziert werden. Diese gehören insbesondere eine Erhöhung der Wassertemperatur seit 1975, zeitigere Eisaufbrüche und eine veränderte Schichtungsstruktur. Dabei wurde im Sommer die Schichtungsstabilität höher und die Intensität der Durchmischung geringer. Das Wasser in der Talsperre erwärmte sich stärker als die Zuflüsse, wodurch diese sich zunehmend tiefer und unterhalb der Thermokline einschichten. Eine zunehmende jährliche Phytoplanktonbiomasse und Artenwechsel innerhalb der Planktongemeinschaft sind weitere Indikatoren für Klimafolgen. Die Kieselalge Aulacoseira subarctica erlangte in Jahren mit warmen Wintern und zeitigen Eisaufbrüchen eine Dominanz während der Frühjahrsentwicklung. Das ungewöhnliche Auftreten dieser Art in aphotischen Tiefen konnte durch eine leichtere Resuspension im Vergleich zu anderen Kieselalgen erklärt werden. Durch zeitige Resuspension gleich nach Eisaufbruch kann A. subarctica ein hohes Inokulum etablieren. Trotz reduzierter Nährstoffzufuhr ins Epilimnion wurde im Sommer die Kieselalge Fragilaria crotonensis vermehrt durch Cyanobakterien verdrängt, was jedoch gut mit den Änderungen der hydrophysikalischen Struktur erklärt werden kann. Obwohl seit 1990 insgesamt mehr Phytoplankton zur Verfügung steht konnte das Crustaceen- Zooplankton in der Talsperre Saidenbach nicht von diesen verbesserten Nahrungsbedingungen profitieren. Die Daphnia-Abundanzen waren seit 1990 stark verringert. Es konnte gezeigt werden, dass der Einfluss des Fischbesatzes unterschätzt wurde und die künstlich ins System eingebrachten Silberkarpfen zeitweise bis zu 70% der zooplanktivoren Fischbiomasse ausgemacht haben. Zoo- planktivore Fische haben einen temperatur- und dichteabhängigen Effekt auf das Zooplankton. Das schnellere Populationswachstum von Daphnia kann bei hohem Besatz mit zooplanktivoren Fischen nicht die erhöhte Fraßaktivität der Fische ausgleichen. Dennoch konnte Temperatur als der wichtigste Einflussfaktor auf die Phänologie des Zooplanktons identifiziert werden, gefolgt von zooplanktivorem Fisch und Nährstoffzufuhr. Die Wichtigkeit von submersen Makrophyten in Flachseen ist gut untersucht. Um den Einfluss von Makropyhten auf die Wasserqualität während des Klimawandels auch in tiefen Seen abschätzen zu können, wurde ein komplexes Seemodell entwickelt. Die Simulationen zeigen, dass sich Makrophyten hauptsächlich positiv auf die Wassergüte auswirken und zur Reduktion von Phytoplankton, insbesondere von Cyanobakterien im Sommer, beitragen. In einem 11 m tiefen See betrug die Reduktion 50 %, in 100 m tiefen oligotrophen Seen immer noch 15 %. Die Konkurrenz um Nährstoffe mit dem Phytoplankton war dabei die ursächliche Makrophyten-Plankton-Interaktion. Submerse Makrophyten könnten für die Restaurierung von tiefen Seen folglich genauso wichtig sein, wie für Flachseen. Das komplette Seemodell enthält hydrophysikalische und ökologische Module und ermöglicht damit weitere umfassende Simulationen zur Untersuchung der Auswirkungen des Klimawandels und zur Evaluation von Adaptionsstrategien für Seen und Talsperren.

Die Gattung Daphnia nimmt eine zentrale Position im pelagischen Nahrungsnetz vieler Standgewässern (Seen, Talsperren) ein. Aufgrund hoher Filtrierleistungen sind Daphnien für das integrierte Gewässermanagement, z.B. im Rahmen einer Nahrungskettenmanipulation (Biomanipulation), von großem Interesse. Ihre relativ einfache Kultivierbarkeit machte sie außerdem zu einen weit verbreiteten Modellorganismus limnologischer Grundlagenforschung. Das Ziel dieser Arbeit besteht darin, ein umfassendes Modellsystem für die Gattung Daphnia zu entwickeln, das wesentliche Kenntnisse der Ökologie von Daphnia miteinander verknüpft, um eine Vernetzung des Wissens zu realisieren. Bestehende Modellansätze konnten diese Aufgabe nur unzureichend erfüllen, weil sie in der Regel einem "Problem-orientierten Ansatz" folgen und somit sehr unterschiedliche inhaltliche Ausrichtungen und verschiedene Modellstrukturen aufweisen. Eine Kopplung bzw. ein direkter Vergleich dieser Modelle ist deshalb in der Regel schwierig. Dieses Modellsystem wurde entwickelt, um ein Instrument zur Synthese des vorhandenen Prozesswissens bereitzustellen, und nicht, um ein spezifisches, abgegrenztes Problem zu bearbeiten; es steht somit diametral zu existierenden Ansätzen. Aus diesem Grunde wird der gewählte Ansatz als ein "Spezies-orientierter Ansatz" bezeichnet. Drei Eigenschaften charakterisieren diesen Ansatz. (i) Das Modellsystem umfasst mehrere hierarchische Ebenen biologischer Organisation, wobei der Schwerpunkt auf den Ebenen des Individuums und der Population liegt. Interaktionen zwischen diesen Organisationsebenen sind ausgeprägt und in Feld- und Laborstudien gut belegt, in existierenden Modellen aber noch unzureichend miteinander verknüpft. (ii) Da eine quantitative Interpretation der Ergebnisse beabsichtigt ist, wurden die Modellausgaben einer umfangreichen Validation an unabhängigen und bereits publizierten Daten unterzogen. (iii) Das Modellsystem zeichnet sich durch einen geschachtelten Aufbau aus (nested design), wodurch eine einfache Erweiterbarkeit des Modells oder auch die separate Anwendung bestimmter Modellkomponenten gewährleistet ist. / The scope of this thesis was to develop a comprehensive model system of the genus Daphnia, a key organism in the pelagic food web of lakes and reservoirs and a widely used model organism in experimental and theoretical ecology. Although its central role in applied and basic research in aquatic ecology is obvious, there are still fundamental problems in modelling the observed dynamics of Daphnia (for details see chapter 2). Therefore, a basic motivation of this work was to use scientific results obtained in independently conducted research for developing a model that brings these results into context. Instead of following a "problem-oriented" paradigm applicable to a single, well defined problem or scientific hypothesis, the underlying concept of the emerging model system was considered to be "species-oriented". Thus, various relevant processes are included into the framework in order to simulate the dynamics of daphnids displayed on different levels of biological organization. To facilitate its application to various problems in ecological research on the genus Daphnia, the model system fulfills the following three important properties: (1) model outputs are thoroughly validated on experimental data in order to guarantee sound quantitative outputs of the model system (2) the system spans over different levels of biological organization with special emphasis laid upon the individual level and the population level (3) the model´s architecture follows a nested design with a defined individual level model that is integrated into a population level model The whole model system is able to describe an individual´s development over time on basis of physiological properties of the organism and, furthermore, how these individual level processes interact with the dynamics on the population level. Due to its nested design, applications of separate submodels (e.g. the individual-level model) are possible.

Indiana University-Purdue University Indianapolis (IUPUI) / Recently there has been a lot of effort in developing systems for sampling and automatically classifying plankton from the oceans. Existing methods assume the specimens have already been precisely segmented, or aim at analyzing images containing single specimen (extraction of their features and/or recognition of specimens as single targets in-focus in small images). The resolution in the existing systems is limiting. Our goal is to develop automated, very high resolution image sensing of critically important, yet under-sampled, components of the planktonic community by addressing both the physical sensing system (e.g. camera, lighting, depth of field), as well as crucial image extraction and recognition routines. The objective of this thesis is to develop a framework that aims at (i) the detection and segmentation of all organisms of interest automatically, directly from the raw data, while filtering out the noise and out-of-focus instances, (ii) extract the best features from images and (iii) identify and classify the plankton species. Our approach focusses on utilizing the full computational power of a multicore system by implementing a parallel programming approach that can process large volumes of high resolution plankton images obtained from our newly designed imaging system (In Situ Ichthyoplankton Imaging System (ISIIS)). We compare some of the widely used segmentation methods with emphasis on accuracy and speed to find the one that works best on our data. We design a robust, scalable, fully automated system for high-throughput processing of the ISIIS imagery.

Consumer-resource interactions are the basic building blocks of every food web. In spite of being a central research theme of longstanding interest in ecology, the mechanisms governing the stability and persistence of consumer-resource interactions are still not entirely understood. In particular, theoretical predictions on consumer-resource stability along gradients of temperature and nutrient enrichment diverge widely and are sometimes in conflict with empirical results. In this thesis I address these issues from the angle of the functional response, which describes a consumer’s feeding rate as a function of resource density. Specifically, I explore mechanistic, nutrient-based consumer-resource interaction models with respect to the influence of feeding behavior (the shape of the functional response), environmental temperature, nutrient enrichment, and resource quality on consumer-resource stability and persistence. In order to parameterize these models I performed extensive laboratory experiments with pairs of freshwater pelagic algae and grazers of the genus Daphnia, which are widespread, ecologically important model organisms. I found a sigmoidal type III functional response in every studied Daphnia-algae species pair. The exact form of its shape is described by an exponent b which is determined by fitting functional response models to the experimental data. A high value of b can stabilize consumer-resource systems under the otherwise destabilizing influence of nutrient enrichment, as predicted by a novel stability criterion relating b to the consumer’s prey handling time, food conversion efficiency and mortality. Estimated parameter values and, consequently, stability predictions are sensitive to the method of parameter estimation, and I propose a new estimation procedure that minimizes parameter uncertainty. Because many consumers’ feeding rates depend on temperature, warming is expected to strongly affect food web stability. In functional response experiments over a broad temperature gradient, I found that the attack rate coefficient and the maximum ingestion rate of Daphnia are hump-shaped functions of temperature. Moreover, the functional response exponent increases with warming towards stronger type III responses. Plugging these findings into a nutrient-based consumer-resource model, I found that predator persistence is a U-shaped function of temperature in nutrient enrichment-temperature space. Enrichment easily turns the system unstable when the consumer has a type II response, whereas a type III response opens up a large region of stability at intermediate, for the consumer optimal, temperatures. These findings reconcile seemingly conflicting results of earlier studies of temperature effects on consumer-resource dynamics, which can be mapped as special cases onto the enrichment-temperature space. I finally demonstrate the utility of three key model ingredients - temperature dependence of rate parameters, a mechanistic description of the dynamics of algal resources, and a type III functional response in Daphnia - by successfully implementing them in the description and explanation of phytoplankton-Daphnia dynamics in a mesocosm experiment exploring effects of warming on the spring succession of the plankton.

Os ecossistemas aquáticos têm sido afetados pelo uso intensivo de agrotóxicos e pela introdução de espécies exóticas. Para melhor avaliar os efeitos diretos e indiretos dessas atividades sobre as comunidades aquáticas, é essencial a utilização de experimentos em escala mais próxima a real. O objetivo desta pesquisa foi analisar os efeitos do agrotóxico Vertimec® 18 CE, cujo princípio ativo é a abamectina, sobre a comunidade planctônica através de um experimento em mesocosmos com duração de 20 dias. Para reproduzir relações biológicas mais complexas e analisar o impacto da introdução de uma espécie exótica sobre o plâncton, inseriram-se girinos de Lithobates catesbeianus em alguns tratamentos. Foram consideradas duas concentrações do agrotóxico (2 μL L-1 em dose múltipla e 8 μL L-1 em dose única) na ausência e presença de girinos, além de dois tratamentos não contaminados (controle com e sem girinos). Variáveis físicas, químicas e biológicas (clorofila a, aspectos qualitativos e quantitativos do fitoplâncton e do zooplâncton) da água e do sedimento foram monitoradas. A partir dos resultados obtidos, verificou-se que os girinos causaram um aumento na concentração de nutrientes na água e uma diminuição do oxigênio dissolvido e da densidade planctônica. Nos tratamentos contaminados, como um efeito indireto do incremento da concentração de nutrientes e da mortalidade das espécies de Cladocera e Copepoda, a densidade do fitoplâncton e de algumas espécies tolerantes de Rotifera aumentou. No tratamento com a concentração mais elevada do agrotóxico, registrou-se a morte de todos os girinos, proporcionando um aumento na concentração de nutrientes na água e na densidade de algumas espécies fitoplanctônicas, que alcançaram uma densidade total de 96.919 ind mL-1. Estes resultados indicam que a inserção dos girinos e a adição do agrotóxico causaram efeitos diretos e indiretos sobre a comunidade planctônica e a qualidade da água. Conclui-se que os mesocosmos podem contribuir para a compreensão de tais efeitos e oferecer subsídios ao manejo de ecossistemas de água doce impactados por atividades antrópicas. / Aquatic ecosystems have been impacted due to the intensive use of pesticides and the introduction of exotic species. It is essential to use more realistic experiments for assessing direct and indirect effects of these activities on aquatic communities. This thesis aimed to analyze the effects of the pesticide Vertimec® 18 EC, whose active ingredient is abamectin, on plankton community through a mesocosm experiment during 20 days. Tadpoles of the species Lithobates catesbeianus were introduced in some treatments to simulate complex biological relationships and to evaluate the impact of the introduction of such exotic species on plankton. Different treatments were analyzed, including one case of no contamination (control) and two cases of contamination: 2 μL L-1 (multiple dose) and 8 μL L-1 (single dose). Such conditions were considered both with absence and presence of tadpoles. Physical, chemical and biological (chlorophyll a, qualitative and quantitative aspects of phytoplankton and zooplankton) variables of water and sediment were also monitored. The tadpoles caused an increase in the concentration of nutrients and a decrease of dissolved oxygen concentration and plankton density. In the contaminated treatments, as an indirect effect of the nutrient increase and the mortality of species of Cladocera and Copepoda, the density of phytoplankton and some tolerant species of rotifers increased. In the treatment with the highest concentration of the pesticide, all the tadpoles died. As a result of the death of the tadpoles, there was nutrient enrichment in the water, with an associated increase in the density of some phytoplankton species, reaching a total density of 96,919 ind mL-1. These results suggested that direct and indirect effects occurred on plankton community and water quality through the addition of the contaminant and the tadpoles. We conclude that the mesocosms might contribute to the understanding of these effects and provide subsidies for management of freshwater ecosystems impacted by human activities.

O estudo de plancton e importante para estimar a biodiversidade em ambientes marinhos, auxiliando na compreensao de sua dinamica e do impacto da atividade humana. Neste trabalho, estudamos as etapas necessarias para a observacao de organismos planctonicos utilizando um sistema holografico digital em linha. A holografia digital permite que as imagens sejam numericamente focalizadas apos sua aquisicao atraves de algoritmos de reconstrucao holografica, o que remove a limitacao da baixa profundidade de foco presente em metodos mais tradicionais, e permite a observacao de um volume maior de agua. E feita uma analise das limitacoes teoricas deste sistema holografico em termos das frequencias presentes no holograma e na sua reconstrucao, observando-se as consequencias para a escolha e implementacao dos algoritmos de reconstrucao. Tambem sao considerados alguns detalhes de implementacao que decorrem da natureza finita e discreta dos algoritmos. No processamento dos hologramas, os organismos sao automaticamente localizados segmentando-se regioes de interesse por meio de uma reconstrucao volumetrica. Estas regioes sao reconstruidas a diversas distancias e e feita a escolha de um plano focal utilizando uma medida de foco. Diversas medidas sao comparadas em imagens obtidas experimentalmente para determinar a mais adequada na focalizacao de organismos planctonicos. Por fim, descrevemos a aplicacao web desenvolvida para o processamento distribuido de imagens holograficas digitais de plancton. / The study of plankton is important for estimating biodiversity in marine environments, helping in the understanding of its dynamics and the impact of human activity. In this work, we study the necessary steps for the observation of planktonic organisms using an inline digital holographic system. Digital holography allows images to be numerically focused after its acquisition through holographic reconstruction algorithms, which removes the limitation of low depth of focus present in more traditional methods, and allows the observation of a bigger volume of water. We present an analysis of the theoretical limitations of this holographic system concerning the frequencies present in the hologram and its reconstruction, noting the consequences for the choice and implementation of reconstruction algorithms. Also considered are some implementation details that stem from the finite and discrete nature of the algorithms. In the processing of holograms, organisms are automatically located by segmenting regions of interest in a volumetric reconstruction. These regions are reconstructed at numerous distances and the choice of a focal plane is made using the values of a focus measure. Various measures are compared in experimentally obtained images to determine the most adequate for focusing planktonic organisms. Finally, we describe a web application developed for distributed processing of digital holographic images of plankton.