<p>The aim of this thesis was to identify general structuring mechanisms in benthic food webs within the framework of ecological stoichiometry theory. Ecological stoichiometry is defined as the balance of multiple chemical substances in ecological interactions and explicitly considers the combined dynamics of key elements such as carbon (C), nitrogen (N) and phosphorus (P). Ecological stoichiometry theory was developed for pelagic environments, thus it must be tested whether the same mechanisms are applicable to benthic environments. </p><p>In this thesis, ecological stoichiometry theory was used as a framework to investigate nutrient pathways in benthic littoral ecosystems. I conducted one invertebrate field sampling and six experiments. In the experiments, factors such as grazing, light, nutrients and fish presence were manipulated.</p><p>The results showed that stoichiometric variability in consumers could mostly be explained by taxa. However, there was some stoichiometric variability due to sampling season, site, and nutrient enrichment. </p><p>Grazing mostly increased periphyton N and P content, although nutrient recycling effects were dependent on grazer stoichiometry. Grazing changed benthic algal community composition by increasing the proportion of grazing resistant algae species. Additionally, grazing decreased algal diversity, especially under nutrient poor conditions. The manipulation of fish presence revealed that fish affected primary producer biomass and stoichiometry through nutrient recycling. </p><p>The manipulation of abiotic factors, such as light and nutrient addition could affect periphyton nutrient content, biomass and benthic algal chlorophyll a content. The separate addition of N or P led to an increase of the added nutrient in the periphyton. Increased light intensities led to a decreased cellular chlorophyll a content and increased C:nutrient ratios. </p><p>This thesis arrives at the conclusion that periphyton-grazer-predator interactions in the benthic are bound by stoichiometric constraints. Nutrient recycling by benthic invertebrates and fish are important mechanism in benthic littoral ecosystems.</p>

<p>Poor food quality can have large negative effects on zooplankton growth and this can also affect food web interactions. The main aims of this thesis were to study the importance of different food quality aspects in <i>Daphnia</i>, to identify potentially important differences among zooplankton taxa, and to put food quality research into a natural context by identifying the importance of food quality and quantity in lakes of different nutrient content.</p><p>In the first experiment, the RNA:DNA ratio was positively related to the somatic growth rate of <i>Daphnia</i>, supporting a connection between P content, RNA content, and growth rate. The second experiment showed that EPA was important for <i>Daphnia</i> somatic growth, and 0.9 µg EPA mg C^-1 was identified as the threshold below which negative effects on <i>Daphnia</i> growth occurred.</p><p>A field survey identified patterns in the PUFA content of zooplankton that could be explained by taxonomy and trophic position. <i>Cladocera</i> enriched EPA and ARA relative to seston, and <i>Copepoda</i> primarily enriched DHA. In a whole-lake experiment, gentle fertilization of an oligotrophicated reservoir increased the seston P content and the biomass of high quality phytoplankton (<i>Cryptophyceae</i>, high EPA content). This was followed by increases in zooplankton and fish biomasses.</p><p>An empirical model based on data from a literature survey predicted that food quantity is most important for zooplankton growth in oligotrophic lakes, and that food quality factors are more important in eutrophic lakes. Thus, zooplankton growth, and energy transfer efficiency in the food web, is predicted to be highest in mesotrophic lakes. The results predict that the strength and nature of food quantity and quality limitation of <i>Daphnia</i> growth varies with lake trophic state, and that some combination of food quantity and/or quality limitation should be expected in nearly all lakes.</p>

Peatlands are peat forming ecosystems in which not fully decomposed plant material builds up the soil. The sequestration of carbon into peat is mainly associated with the bryophyte genus Sphagnum (peat mosses), which dominate and literally form most peatlands. The responses of Sphagnum to environmental change help us to understand peatland development and function and to predict future changes in a rapidly changing world. In this thesis, the overarching aim was to use ecophysiological traits to investigate mechanisms behind the response of Sphagnum to elevated N deposition, and, processes connected to ecosystem shift and ecosystem function of peatlands. Regarding elevated N deposition, three experiments were performed at different scales (country-wide to greenhouse). Independent of scale and species, apical tissue N concentration increased with increasing N input until N saturation was reached. Maximum photosynthetic rate, a trait evaluating photosynthetic capacity, increased with N input and could be well predicted by tissue N concentration. Thus, the physiological responses of Sphagnum to N deposition are often positive and I found no evidence of toxic effects. Production did, however, not increase with N input, and results of the N:P ratio suggested that P limitation, and possibly other elements, might hamper growth under high N input. The effect of P limitation was, in contrast to current view, most pronounced in fast growing species indicating species specific responses to nutrient imbalance. I explored the puzzling, but historically frequently occurring, rich fen to bog ecosystem shift; a shift from a species-rich ecosystem dominated by brown mosses, to a species-poor one with greater carbon storage that is Sphagnum-dominated. The bog-dwelling species of Sphagnum grew well, to our surprise, when in contact with rich fen water but was not a strong competitor compared to rich fen Sphagnum species. If submerged under rich fen water (high pH), the bog Sphagnum species died while rich fen species of Sphagnum were unaffected. These results show that differences in two physiological traits (growth rate and tolerance to flooding) among species, can explain when a peatland ecosystem shift might occur. In the last study, the function of peatlands was related to trade-offs between traits and allometric scaling in Sphagnum. Results suggested that growth strategies are determined by the distribution of Sphagnum relative to the water table in order to minimize periods with suboptimal hydration. Allometric analyses stressed the importance of resource allocation among and within shoots (apical part vs. stem), although the allocation patterns in Sphagnum were not always consistent with those of vascular plants. Interestingly, data indicated a trade-off between photosynthetic rate and decomposition rate among Sphagnum species.

L'objectiu d'aquest estudi és el d'investigar sobre l'ús de matèria orgànica per part dels fongs i bacteris que colonitzen diferents substrats bentònics en rius Mediterranis i analitzar l'efecte dels factors ambientals i antròpics sobre l'estabilitat estructural i funcional de les comunitats del biofilm. La metodologia emprada en aquest estudi consisteix en: i) anàlisi de la biomassa bacteriana i fúngica, ii) anàlisi de la composició de les comunitats bentòniques (identificació d'hifomicets aquàtics i anàlisi del 16S rDNA bacterià), i iii) anàlisi de l'activitat enzimàtica extracel·lular relacionada amb el reciclatge de matèria orgànica en rius. / This study aimed to investigate on the use of organic matter by fungi and bacteria inhabiting different benthic substrata and to analyze the effect of environmental and anthropogenic perturbations on the structural and functional stability of biofilms. The following methodologies has been used in this study: i) analysis of fungal and bacterial biomass, ii) analysis of benthic community composition (identification of hyphomycete taxa, analysis of the bacterial 16S rDNA), and iii) analysis of extracellular enzyme activities involved in organic matter recycling in rivers.

Amb aquest treball es preté comprovar la universalitat de la relació estequiomètrica de Redfield i recercar patrons consistents de les desviacions d'aquesta raó promig en ambients oligotrofics. Per tal de dur a terme aquest propòsit s'han desenvolupat relacions estequiomètriques en el compartiment particulat i dissolt orgànic i inorgànic per el C, N, P i Si.La estequiometria dels nutrients a l'oceà s'ha examinat al Mar Mediterrani i a l'Oceà Atlàntic subtropical, mentre que la zona costenera del Mar Mediterrani ha servit per estudiar aquestes relacions sota l'influència d'aports terrestres. En el segon capítol de la tesi es descriuen patrons meridionals del fluxe de nitrogen i fósfor vers la capa biogènica de l'Oceà Atlàntic Central. La raó promig entre el fluxe difusiu del nitrogen inorgànic dissolt (DIN) i del fósfor (DIP) es mostra similar a la raó de Redfield al llarg de l'Oceà Atlàntic Central, però tendeix a valors per sota dels establerts per Redfield a la part sud del trajecte i superiors a la raó de Redfield al centre del gir sudtropical. La raó N:P del fluxe difusiu i la raó N:P dels nutrients dissolts inorgànics en la capa biogènica es troben fortament correlacionats, mentre que no existeix cap correlació amb els valors de la raó N:P de les aigües intermitges. Els resultats trobats en aquest capítol de la tesi indiquen que la recirculació vertical de nutrients a la capa biogènica de l'oceà Atlàntic Central és capaç d'operar amb raons estequimètriques que difereixen de Redfield i per tant els components biogènics i biolítics s'adapten a les variacions locals de la raó de Redfield. La hipòtesis que existeixen raons estequiomètriques previsibles en la reserva oceànica de material dissolt orgànic que es troben en equilibri amb la reserva del material particulat orgànic i dissolt inorgànic, es corrobora en el tercer capítol d'aquesta tesi. La majoria del carboni orgànic present en aigües oligotrofiques del Mediterrani estratificat es troba en forma d'orgànic dissolt, mentre que el POC (carboni orgànic particulat) representa un percentatge menor. El nitrogen i fósfor orgànic dissolt que comprenen el 50-80% del "pool" total de P i N a la capa biogènica, decreix en percentarge a la capa biolítica. S'ha comprovat una distribució uniforme del nitrogen disolt total (TDN). L'increment en el percentatge de N inorgànic disolt i el decreixement en percentatge de N orgànic dissolt amb la fondària, és un indicador clar de l'equilibri dinàmic que existeix entre les reaccions bioquímiques entre les reserves oceàniques. Mitjançant un sistema de balanços, s'estableix un intercanvi de nutrients (exportació-importació) entre els pools dissolt orgànic i dissolt inorgànic en el sistema. S'ha comprovat que el fluxe de nitrogen orgànic dissolt (DON) excedeix al fluxe difusiu de nitrogen inorgànic i per tant els aports atmosfèrics i terrestres són importants en aquesta regió.En el quart capítol de la tesi es descriu el lligam existent entre el component pelàgic i els component del bentos d'una àrea litoral del Mar Mediterrani, és a dir un compartiment anabolic que produeix matèria orgànica i un de catabolic que actúa com agent oxidant de la reserva de matèria orgànica del sistema. El compartiment pelàgic es mostra heterotrofic. Al mateix temps, existeix una contribució important de material terrestre als sediments. Per contra, el compartiment bentònic és autotrofic on el dèficit en la producció grossa es compensa amb l'excés de producció neta del sistema. Mitjançant la quantificació simultànea dels fluxes anuals de sedimentació per al C, N, P i Si així com dels fluxes sediment-aigua de les espècies orgàniques i inorgàniques dissoltes, s'ha establert un sistema de balanços de matèria del sistema. El compartiment bentonic es configura com a sumider o exportador de matèria orgànica, degut als aports terrestres de carboni en el sistema.Els patrons de distribució de nutrients derivats dels resultats dels capítols anteriors es comproven mitjançant un experiment d'addició de nutrients. En el capítol cinquè d'aquesta tesi s'estudien els canvis en la distribució de nutrients en les diferents reserves nutricionals d'un sistema quan es troba sotmès a aports controlats de nutrients. S'ha comprovat que, mentre el tamany relatiu de la reserva de nutrients inorgànics dissolts no varia amb l'increment de nutrients en el medi. Hi ha una tendència a l'increment del tamany relatiu de la reserva del material particulat, paral.lela a un decreixement simultani de la reserva de material dissolt orgànic. Aquest experiment contribueix a verificar el paper del DOM (matèria dissolta orgànica) com a principal reserva nutricional en sistemes oliogotròfics. / The stoichiometric ratios are powerful tools to model basic biogeochemical patterns of the sea when the fluxes of a single element are known. It is, therefore essential to understand the full implications of variable stoichiometries to predict the effect of the living components of the ocean on biogeochemical processes. Here, the stoichiometry between C, N, P and Si in different chemical pools (particulate and dissolved organic and dissolved inorganic matter) were examined in contrasting oceanic and littoral ecosystems, and the changes in nutrient partitioning in response to nutrient inputs was tested through experimental research. In the Central Atlantic Ocean, the average ratio between dissolved inorganic nitrogen and phosphorus in the estimated vertical diffusive fluxes was similar to the Redfield ratio, but tended to be above the Redfield ratio at the center of the South subtropical gyre. The N:P supply ratio and the N:P ratio of dissolved inorganic nutrients in the biogenic layer were strongly correlated, but were not positively correlated to that in the intermediate waters. The vertical nutrient conveyor belt of nutrients in the upper waters operates relatively independently of the underlying waters in the Central Atlantic, so that both the biogenic and the biolythic components should be able to adapt to local variation about the Redfield ratio.In the stratified NW Mediterranean Sea, the stoichiometry between dissolved inorganic, organic and particulate organic matter pools indicated an excess nitrogen relative to phosphorus, particularly in the biolythic layer, as well as a general silicate deficiency relative to both N and P. Most (> 80 %) of the organic carbon was present as dissolved organic carbon, with POC representing a minor percent of total organic C throughout the water column. The increasing C/N ratio of DOM with depth indicates that N is recycled faster than C in the DOM. There exists a dynamic equilibrium between the biological transformations between these pools with depth, with a dominance of DON production in surface waters and remineralization in the underlying layers, from which dissolved inorganic nitrogen is re-supplied to the biogenic layer. Alocthonous N inputs must be important in the region since the downward DON flux exceeded the diffusive DIN supply. The coupling between anabolic and catabolic compartments of a littoral area in the NW Mediterranean Sea are characterized. The pelagic compartment was heterotrophic, supported by significant allochthonous inputs of land material, whereas the benthic compartment was autotrophic, with the excess net benthic community production balancing the deficit in pelagic community production, leading to a system in metabolic equilibrium. Sedimentary inputs of phosphorus and silicon were compensated by sediment release of phosphate and silicate, whereas nitrogen was lost or accumulated in the sediment compartment. Carbon inputs to the benthic compartment also exceeded requirements, due to the allocthonous subsidies to the system, so that the benthic compartment stored or exported organic carbon.Experimental nutrient additions lead to a parabolic change in C/N and C/P ratios in the dissolved organic matter with increasing nutrient inputs. The relative size of the dissolved inorganic nutrient pools did not vary, but there was a tendency towards an increase in the relative size of the particulate pool at the expense of a decrease in the relative importance of DOM as a reservoir of N, P and C, with increasing nutrient inputs.The relative importance of different nitrogen pools was examined in relation to the total nutrient stoichiometry of the oligotrophic system. The ratio of dissolved inorganic nutrients reported in the research presented is indicative of a general deficiency in nitrogen compared to the global ratios reported in literature. The dissolved organic matter was highly depleted in P relative to N and C at all locations investigated and the resulting POC:PON ratio (11.7) of this study in the particulate pool deviates greatly from the literature values which approximates Redfield value (5.5-6). The shift of the dominance of DON towards PON at TOC/TN values higher than 20 on the oligothrophic areas of the study, gives evidence of increasing carbon export fluxes in a system dominated by particulate pool and points to the effect of the biota on the gradient-driven export to sinking carbon fluxes in the ocean.

The influenza virus hemagglutinin (HA) surface protein is a primary antigenic target for neutralization of viral infection. HA also mediates membrane fusion between the virus and a cell, which is the first critical step during infection. Traditional techniques to study infection neutralization by antibodies or the membrane fusion process rely on ensemble measurements, confounding the precise mechanism of infection neutralization and obscuring transient conformational intermediates. This dissertation describes advances made in a fluorescence microscopy-based single-particle fusion assay to overcome the limitations of ensemble measurements in these types of studies. Virus particles are labeled to visualize lipid mixing between a virus and a target membrane formed upon a glass or polymer support. Optionally, the viral lumen can be labeled to visualize the subsequent release of viral contents.

A new combustion zone analytical model has been developed in which the combustion front temperature may be calculated. The model describes in the combustion zone, the amount of fuel burned based on reaction kinetics, the fuel concentration and produced gas composition based on combustion stoichiometry, and the amount of heat generated based on a heat balance. Six runs were performed in a 3-inch diameter, 40-inch long steel combustion tube with Jobo crude oil (9-11°API) from the Orinoco Belt in Venezuela. These runs were carried out with air containing three values of oxygen concentration, 21%, 30%, and 40%. The weight percentage of sand, clay, water, and oil in the sand mix was kept constant in all runs at 86.6%, 4.7%, 4.0%, and 4.7% respectively. Injection air rates (3 L/min) as well as the production pressure (300 psig) were kept constant in all runs. The results indicate that the calculated combustion zone temperatures and temperature profiles are in good agreement with the experimental data, for the range of oxygen concentration in the injected air. The use of oxygen-enriched air slightly increased the combustion front temperature from 440°C in a 21 mole % O2 concentration to a maximum of 475°C for air with 40 mole % O2 concentration. Oxygen-enriched air injection also increased the combustion front velocity from 13.4 cm/hr (for 21% oxygen) to 24.7 cm/hr (for 40% oxygen), thus reducing the start of oil production from 3.3 hours (for 21% oxygen) to 1.8 hours (for 40% oxygen). In the field, the use of oxygen-enriched air injection could translate into earlier oil production compared to with not-enriched air injection. The new analytical model for the combustion zone developed in this study will be beneficial to future researchers in understanding the effect of oxygen-enriched in-situ combustion and its implications on the combustion front temperature and combustion front thickness.

No description available.

Competitive-Consecutive and Competitive-Parallel reactions are both mixing sensitive reactions; the yield of desired product from these reactions depends on how fast the reactants are brought together. Recent experimental results have suggested that the mixing effect may depend strongly on the stoichiometry of the reactions. To investigate this, a 1-D, non-dimensional, reaction-diffusion model at the micro-mixing scale has been developed. Assuming constant mass concentration and diffusivities, systems of PDEs have been derived on a mass fraction basis for both types of reactions. A single general Damkhler number and specific dimensionless reaction rate ratios were derived for both reaction schemes. The resulting dimensionless equations were simulated to investigate the effects of mixing, reaction rate ratio and stoichiometry of the reactions. It was found that decreasing the striation thickness and the dimensionless rate ratio maximizes yield for both types of reactions and that the stoichiometry has a considerable effect on yield. All three variables were found to interact strongly. Phase plots showing the interactions between the three variables were developed.

The aim of this thesis was to identify general structuring mechanisms in benthic food webs within the framework of ecological stoichiometry theory. Ecological stoichiometry is defined as the balance of multiple chemical substances in ecological interactions and explicitly considers the combined dynamics of key elements such as carbon (C), nitrogen (N) and phosphorus (P). Ecological stoichiometry theory was developed for pelagic environments, thus it must be tested whether the same mechanisms are applicable to benthic environments. In this thesis, ecological stoichiometry theory was used as a framework to investigate nutrient pathways in benthic littoral ecosystems. I conducted one invertebrate field sampling and six experiments. In the experiments, factors such as grazing, light, nutrients and fish presence were manipulated. The results showed that stoichiometric variability in consumers could mostly be explained by taxa. However, there was some stoichiometric variability due to sampling season, site, and nutrient enrichment. Grazing mostly increased periphyton N and P content, although nutrient recycling effects were dependent on grazer stoichiometry. Grazing changed benthic algal community composition by increasing the proportion of grazing resistant algae species. Additionally, grazing decreased algal diversity, especially under nutrient poor conditions. The manipulation of fish presence revealed that fish affected primary producer biomass and stoichiometry through nutrient recycling. The manipulation of abiotic factors, such as light and nutrient addition could affect periphyton nutrient content, biomass and benthic algal chlorophyll a content. The separate addition of N or P led to an increase of the added nutrient in the periphyton. Increased light intensities led to a decreased cellular chlorophyll a content and increased C:nutrient ratios. This thesis arrives at the conclusion that periphyton-grazer-predator interactions in the benthic are bound by stoichiometric constraints. Nutrient recycling by benthic invertebrates and fish are important mechanism in benthic littoral ecosystems.