Thesis (Ph. D.)--Uppsala University, Dept. of Physical Geography, Division of Hydrology, 1990. / Cover title. Errata sheet inserted.

Thesis (Ph. D.)--Oregon State University, 2007. / Printout. Includes bibliographical references. Also available on the World Wide Web.

Nyanza Gulf is a large, shallow and long river-influenced embayment located in northeastern Lake Victoria. The gulf opens to the main lake through the narrow and deep Rusinga Channel, the exchange zone between the two ecosystems with different physical chemical and biogeochemical conditions. The main goals of this study are to characterize physicochemical and nutrient gradients along the gulf-main-lake transect, characterize and quantify the water and nutrient fluxes between the gulf and the main lake, and assess the response of phytoplankton community and photosynthesis to the spatially varying physical and nutrient conditions along the study transect. Between March 2005 and March 2006, measurements of physicochemical profiles as well as nutrient and the phytoplankton community analysis were conductued monthly along the study transect. Additionally, analysis of different surficial sediment phosphorus fractions was done in order to asses the potential role of bottom sediment in contributing to phosphorus enrichment in the lake water column. A box mass balance model was used to calculate the exchange of water and nutrient fluxes between different zones along the study transect and to estimate ecosystem metabolism in the gulf and the channel. Spatial variability in physicochemical and biogeochemical conditions was observed along the study transect, especially between the shallow and river-influenced inner-gulf, the deep and physically active Rusinga Channel, and the main lake, mainly in response to river inputs and varying morphometry along the study transect. The gulf had significantly higher electrical conductivity (EC), turbidity, total nitrogen (TN), and dissolved reactive silica (DRSi) but the levels declined monotonically along the channel in response to mixing with the main lake water. The channel and the main lake had, respectively, significantly higher dissolved inorganic nitrogen (DIN) and soluble reactive phosphorus (SRP) compared to the gulf. Spatial variability in morphometry and exposure to varying wind forcing lead to differential mixing and differential heating and cooling along the transect, resulting in density driven fronts and horizontal exchange of water and nutrients between the gulf and the main lake. Upwelling and downwelling maintained mixing conditions in the channel which consequently influenced nutrient recycling, the light environment and hence affecting phytoplankton community composition and productivity. The net residual water flow from the gulf to the main lake was 36 m3/s but the mixing flux was approximately 20 times higher and both fluxes accounted for a gulf exchange time of 1981 days. The advective and mixing fluxes between the gulf and the main lake resulted in net export of dissolved inorganic phosphorus (DIP; 400 kg P/d) from the main lake into the gulf and net export of DRSi (10 t Si/d) from the gulf into the main lake. In the deep, narrow and physically active Rusinga Channel there was net production of dissolved nutrients whereas in the gulf there was net consumption of dissolved nutrients, which helped to maintain high net ecosystem production (NEP; 566 mg C/m2/d) in the gulf in contrast the channel which showed net heterotrophy. The high NEP in the gulf and the associated high nutrient demand coupled with possibly low SRP to DIN supply ratio lead to P limitation of algal growth in the gulf as indicated by all indicators of nutrient status. This has important implications for management since increased P input into the gulf will translate into increased algal blooms in the gulf and therefore compromise water quality. Spatial variability in physical conditions and nutrient status along the study transect influenced phytoplankton community composition and photosynthesis. The shallow and turbid gulf was dominated by cyanobacteria but diatoms dominated in the channel in response to reduced turbidity and increased physical mixing and nutrient availability (DRSi, SRP). In the main lake seasonal stratification and deep mixing depth favoured both cyanobacteria and diatoms. The phytoplankton community in channel had a higher photosynthetic capacity (Fv/Fm, PBm) compared to both the gulf and the main lake.

Nyanza Gulf is a large, shallow and long river-influenced embayment located in northeastern Lake Victoria. The gulf opens to the main lake through the narrow and deep Rusinga Channel, the exchange zone between the two ecosystems with different physical chemical and biogeochemical conditions. The main goals of this study are to characterize physicochemical and nutrient gradients along the gulf-main-lake transect, characterize and quantify the water and nutrient fluxes between the gulf and the main lake, and assess the response of phytoplankton community and photosynthesis to the spatially varying physical and nutrient conditions along the study transect. Between March 2005 and March 2006, measurements of physicochemical profiles as well as nutrient and the phytoplankton community analysis were conductued monthly along the study transect. Additionally, analysis of different surficial sediment phosphorus fractions was done in order to asses the potential role of bottom sediment in contributing to phosphorus enrichment in the lake water column. A box mass balance model was used to calculate the exchange of water and nutrient fluxes between different zones along the study transect and to estimate ecosystem metabolism in the gulf and the channel. Spatial variability in physicochemical and biogeochemical conditions was observed along the study transect, especially between the shallow and river-influenced inner-gulf, the deep and physically active Rusinga Channel, and the main lake, mainly in response to river inputs and varying morphometry along the study transect. The gulf had significantly higher electrical conductivity (EC), turbidity, total nitrogen (TN), and dissolved reactive silica (DRSi) but the levels declined monotonically along the channel in response to mixing with the main lake water. The channel and the main lake had, respectively, significantly higher dissolved inorganic nitrogen (DIN) and soluble reactive phosphorus (SRP) compared to the gulf. Spatial variability in morphometry and exposure to varying wind forcing lead to differential mixing and differential heating and cooling along the transect, resulting in density driven fronts and horizontal exchange of water and nutrients between the gulf and the main lake. Upwelling and downwelling maintained mixing conditions in the channel which consequently influenced nutrient recycling, the light environment and hence affecting phytoplankton community composition and productivity. The net residual water flow from the gulf to the main lake was 36 m3/s but the mixing flux was approximately 20 times higher and both fluxes accounted for a gulf exchange time of 1981 days. The advective and mixing fluxes between the gulf and the main lake resulted in net export of dissolved inorganic phosphorus (DIP; 400 kg P/d) from the main lake into the gulf and net export of DRSi (10 t Si/d) from the gulf into the main lake. In the deep, narrow and physically active Rusinga Channel there was net production of dissolved nutrients whereas in the gulf there was net consumption of dissolved nutrients, which helped to maintain high net ecosystem production (NEP; 566 mg C/m2/d) in the gulf in contrast the channel which showed net heterotrophy. The high NEP in the gulf and the associated high nutrient demand coupled with possibly low SRP to DIN supply ratio lead to P limitation of algal growth in the gulf as indicated by all indicators of nutrient status. This has important implications for management since increased P input into the gulf will translate into increased algal blooms in the gulf and therefore compromise water quality. Spatial variability in physical conditions and nutrient status along the study transect influenced phytoplankton community composition and photosynthesis. The shallow and turbid gulf was dominated by cyanobacteria but diatoms dominated in the channel in response to reduced turbidity and increased physical mixing and nutrient availability (DRSi, SRP). In the main lake seasonal stratification and deep mixing depth favoured both cyanobacteria and diatoms. The phytoplankton community in channel had a higher photosynthetic capacity (Fv/Fm, PBm) compared to both the gulf and the main lake.

Phytoplankton chlorophyll-a fluorescence, measured in situ, can be applied as a tool to estimate primary production in the ocean over a large range of temporal and spatial scales. This non-invasive technique allows for fast assessments of photo-physiological parameters in contrast to the traditional methodologies (14C uptake and O2 evolution). The main photo-physiological parameters derived by the available instruments are yields, and as such, require careful interpretation. The comprehension of the main sources of variability of the photochemical and the light absorption efficiencies in marine phytoplankton has increased in the past years, largely by studies using monospecific cultures. In natural communities, however, the development of primary production models based on chlorophyll-a fluorescence remain limited as they are simultaneously subjected to a wide range of environmental and biological factors. This study will test the hypothesis that photo-physiological models for primary production estimates can be improved when key phytoplankton features, such as the pigments composition and dominant cell size, are taking into account. The approach was to contrast the photo-physiological parameters derived from measurements in distinct oceanographic regions, as well as those derived in a specific environment with presented different nutrient concentration according to the time of sampling. In addition, we showed for monospecific cultures, how the photo-physiological parameters are quantitatively related to the production of carbon under the interactive effects of taxonomic composition and cell size. The proportions of photosynthetic and photoprotective pigments present in the community were related to the bulk photochemical efficiency and the cross-section of light absorption, but varied among oceanographic regions and the depth of the water column. A parameterization of fluorescence-derived primary production rates, using four dominant size classes, was derived for natural phytoplankton communities under different nutrients conditions in a coastal environment, showing that the parameters differed among size classes above a threshold of nutrient concentration. The direct conversion rates between fluorescence-derived primary production and carbon assimilation rates, computed for two distinct phytoplankton cell sizes grown in controlled laboratorial conditions, showed that cell size strongly influences the efficiency of light absorption and photochemistry, however species-specific responses in photosynthetic energy allocation dominated the differences observed in how absorbed light is utilized to carbon assimilation, i.e., in the electron requirements for carbon assimilation. The results highlighted the importance of the tight coupling of nutrients availability and phytoplankton communities, as well as for measurements of chlorophyll-a fluorescence in the ocean and primary production models. This work presents a novel contribution to the increasing efforts to apply fluorescence-based techniques to understand and parameterize primary production estimates in marine systems, especially at highly dynamic environments. / A fluorescência da clorofila-a do fitoplâncton, medida in situ, pode ser uma ferramenta para estimar produção primária no oceano em grande escala temporal e espacial. Esta técnica não-invasiva permite análises rápidas de parâmetros foto-fisiológicos ao contrário de metodologias tradicionais (assimilação de 14C e produção de O2). Os principais parâmetros foto-fisiológicos de instrumentos disponíveis hoje tratam-se de eficiências, e como tal, requerem cuidados em serem interpretados. A compreensão das principais fontes de variabilidade da eficiência fotoquímica e de absorção de luz no fitoplâncton marinho tem aumentado nos últimos anos, em sua maioria em cultivos monoespecíficos. Em comunidades naturais, entretanto, o desenvolvimento de modelos de produção primária baseados na fluorescência da clorofila-a ainda é limitado uma vez que estão sujeitos à uma ampla gama de fatores ambientais e biológicos. Esse estudo testa a hipótese de que modelos foto-fisiológicos para estimar produção primária podem ser aprimorados considerando-se características fundamentais do fitoplâncton, como a composição de pigmentos e tamanho celular dominante. A estratégia foi contrastar parâmetros foto-fisiológicos derivados de medidas em regiões oceanográficas distintas, assim como medidas em um ambiente específico com diferentes concentrações de nutrientes ao longo do período amostrado. Adicionalmente, apresentamos através de cultivos monoespecíficos, como parâmetros foto-fisiológicos são quantitativamente relacionados à produção de carbono e os efeitos interativos da composição taxonômica e tamanho celular nessa relação. A proporção de pigmentos fotossintéticos e fotoprotetores da comunidade foram relacionados à eficiência fotoquímica e seção transversal de absorção de luz, porém variaram de acordo com a região oceanográfica e profundidade na coluna d\'água. Uma parameterização de taxas de produção primária derivadas da fluorescência, usando quatro classes de tamanho dominantes, foi proposta para comunidades naturais de fitoplâncton sob condições de nutrientes diferentes em um ambiente costeiro, mostrando que os parâmetros diferiram entre as classes de tamanho acima de um limiar de concentração de nutrientes. As taxas de conversão diretas entre produção primária derivada da fluorescência e taxas de assimilação de carbono, computadas para dois tamanhos de fitoplâncton crescidos em condições controladas em laboratório, mostraram que tamanho celular influencia as eficiências de absorção de luz e fotoquímica, porém respostas espécie-específicas na alocação de energia fotossintética dominaram as diferenças observadas em como a luz absorvida é utilizada para assimilação de carbono, ou seja, na razão de elétrons exigidos para assimilação de carbono. Os resultados destacam a importância do acoplamento da disponibilidade de nutrientes com a comunidade fitoplanctônica, assim como das medidas de fluorescência da clorofila-a no oceano e nos modelos de produção primária. Este trabalho apresenta uma contribuição inédita nos esforços crescentes em aplicar técnicas baseadas na fluorescência para entender e parameterizar estimativas de produção primária nos sistemas marinhos, especialmente em ambientes altamente dinâmicos.

Phytoplankton chlorophyll-a fluorescence, measured in situ, can be applied as a tool to estimate primary production in the ocean over a large range of temporal and spatial scales. This non-invasive technique allows for fast assessments of photo-physiological parameters in contrast to the traditional methodologies (14C uptake and O2 evolution). The main photo-physiological parameters derived by the available instruments are yields, and as such, require careful interpretation. The comprehension of the main sources of variability of the photochemical and the light absorption efficiencies in marine phytoplankton has increased in the past years, largely by studies using monospecific cultures. In natural communities, however, the development of primary production models based on chlorophyll-a fluorescence remain limited as they are simultaneously subjected to a wide range of environmental and biological factors. This study will test the hypothesis that photo-physiological models for primary production estimates can be improved when key phytoplankton features, such as the pigments composition and dominant cell size, are taking into account. The approach was to contrast the photo-physiological parameters derived from measurements in distinct oceanographic regions, as well as those derived in a specific environment with presented different nutrient concentration according to the time of sampling. In addition, we showed for monospecific cultures, how the photo-physiological parameters are quantitatively related to the production of carbon under the interactive effects of taxonomic composition and cell size. The proportions of photosynthetic and photoprotective pigments present in the community were related to the bulk photochemical efficiency and the cross-section of light absorption, but varied among oceanographic regions and the depth of the water column. A parameterization of fluorescence-derived primary production rates, using four dominant size classes, was derived for natural phytoplankton communities under different nutrients conditions in a coastal environment, showing that the parameters differed among size classes above a threshold of nutrient concentration. The direct conversion rates between fluorescence-derived primary production and carbon assimilation rates, computed for two distinct phytoplankton cell sizes grown in controlled laboratorial conditions, showed that cell size strongly influences the efficiency of light absorption and photochemistry, however species-specific responses in photosynthetic energy allocation dominated the differences observed in how absorbed light is utilized to carbon assimilation, i.e., in the electron requirements for carbon assimilation. The results highlighted the importance of the tight coupling of nutrients availability and phytoplankton communities, as well as for measurements of chlorophyll-a fluorescence in the ocean and primary production models. This work presents a novel contribution to the increasing efforts to apply fluorescence-based techniques to understand and parameterize primary production estimates in marine systems, especially at highly dynamic environments. / A fluorescência da clorofila-a do fitoplâncton, medida in situ, pode ser uma ferramenta para estimar produção primária no oceano em grande escala temporal e espacial. Esta técnica não-invasiva permite análises rápidas de parâmetros foto-fisiológicos ao contrário de metodologias tradicionais (assimilação de 14C e produção de O2). Os principais parâmetros foto-fisiológicos de instrumentos disponíveis hoje tratam-se de eficiências, e como tal, requerem cuidados em serem interpretados. A compreensão das principais fontes de variabilidade da eficiência fotoquímica e de absorção de luz no fitoplâncton marinho tem aumentado nos últimos anos, em sua maioria em cultivos monoespecíficos. Em comunidades naturais, entretanto, o desenvolvimento de modelos de produção primária baseados na fluorescência da clorofila-a ainda é limitado uma vez que estão sujeitos à uma ampla gama de fatores ambientais e biológicos. Esse estudo testa a hipótese de que modelos foto-fisiológicos para estimar produção primária podem ser aprimorados considerando-se características fundamentais do fitoplâncton, como a composição de pigmentos e tamanho celular dominante. A estratégia foi contrastar parâmetros foto-fisiológicos derivados de medidas em regiões oceanográficas distintas, assim como medidas em um ambiente específico com diferentes concentrações de nutrientes ao longo do período amostrado. Adicionalmente, apresentamos através de cultivos monoespecíficos, como parâmetros foto-fisiológicos são quantitativamente relacionados à produção de carbono e os efeitos interativos da composição taxonômica e tamanho celular nessa relação. A proporção de pigmentos fotossintéticos e fotoprotetores da comunidade foram relacionados à eficiência fotoquímica e seção transversal de absorção de luz, porém variaram de acordo com a região oceanográfica e profundidade na coluna d\'água. Uma parameterização de taxas de produção primária derivadas da fluorescência, usando quatro classes de tamanho dominantes, foi proposta para comunidades naturais de fitoplâncton sob condições de nutrientes diferentes em um ambiente costeiro, mostrando que os parâmetros diferiram entre as classes de tamanho acima de um limiar de concentração de nutrientes. As taxas de conversão diretas entre produção primária derivada da fluorescência e taxas de assimilação de carbono, computadas para dois tamanhos de fitoplâncton crescidos em condições controladas em laboratório, mostraram que tamanho celular influencia as eficiências de absorção de luz e fotoquímica, porém respostas espécie-específicas na alocação de energia fotossintética dominaram as diferenças observadas em como a luz absorvida é utilizada para assimilação de carbono, ou seja, na razão de elétrons exigidos para assimilação de carbono. Os resultados destacam a importância do acoplamento da disponibilidade de nutrientes com a comunidade fitoplanctônica, assim como das medidas de fluorescência da clorofila-a no oceano e nos modelos de produção primária. Este trabalho apresenta uma contribuição inédita nos esforços crescentes em aplicar técnicas baseadas na fluorescência para entender e parameterizar estimativas de produção primária nos sistemas marinhos, especialmente em ambientes altamente dinâmicos.