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Optimering av den kemiska reningen vid Fläskebo deponi / Optimization of the Chemical Treatment at Fläskebo LandfillNilsson, Anna January 2006 (has links)
<p>Landfill leachate contains a variety of contaminants and is created when rainwater percolates the landfill. For landfill management the leachate is the main issue that can cause problems to the environment. At the landfill of Fläskebo, Renova AB treats the leachate in a local treatment plant. The treatment consists of a chemical treatment step with chemical precipitation, flocculation, sedimentation and filtration, and a final step with a carbon and peat filter. Renova has to ensure that the condition of the leachate reaches the regulation set for the landfill before it is released to the recipient. This regulation has not yet been established and a final suggestion will be given to the county administrative board in spring 2006.</p><p>In this master thesis the chemical treatment of Fläskebo is optimized. A comparison between the control program and the regulation was made to estimate the contamination of the leachate. Also the effectiveness of the two steps is evaluated. For optimization, the leachate was first tested in a laboratory with different coagulants and flocculants.</p><p>The purpose was to increase the precipitation of particles and metals; arsenic, cadmium, chromium, mercury, lead, copper, nickel and zinc through sweep floc coagulation and hydroxide precipitation. After the laboratory tests the precipitation was tested in the treatment plant with higher pH and coagulant. Also the process control for sodium hydroxide was examined.</p><p>The leachate had a small content of organic matter and nutrients, but had a large content of halogenated substances (AOX) and the heavy metals nickel and copper. High concentrations of contaminants were reduced better than low concentrations in the two treatment steps. The carbon and peat filter material also caused an increase of the arsenic content in the leachate after filtration. From the laboratory work the results showed a better reduction of metals with iron-chloride, PlusJÄRN and the anjonic polyacrylamid, Fennopol A. Because of the high content of chloride the iron- sulphate, PurFect was chosen for further tests. The optimal pH for the heavy metals arsenic, zinc, copper and nickel was between pH 9 and 9, 5. The precipitation in the treatment plant showed better results with sodium hydroxide and a higher pH, pH 9 in the flocculation basin. An increase of the coagulant PurFect from 202 mg/l to 225 mg/l meant only a higher chemical cost. The process control of sodium hydroxide showed an oscillating and unstable control performance, which may lead to a higher consumption of chemicals.</p> / <p>Lakvatten har varierande föroreningsgrad och karaktär och bildas då regnvatten perkolerar igenom en deponi. För en deponiverksamhet är lakvattnet den huvudsakliga påverkan på den omgivande miljön. Vid en av Renova ABs deponier, Fläskebo, utanför Göteborg sker reningen av vattnet i den lokala reningsanläggningen. Reningen består av kemisk fällning, flockning, sedimentering och filtrering samt ett kol- och torvfilter. För att få släppa ut lakvattnet har Renova just nu ett prövotidsvillkor på lakvattnets kvalité och till våren 2006 skall förslag på slutgiltiga utsläppsvillkor lämnas till Länsstyrelsen.</p><p>I detta examensarbete har den kemiska reningen av lakvattnet från Fläskebo optimerats utifrån lakvattnets karaktär. Lakvattnets föroreningsgrad bedömdes efter en jämförelse av analysresultatet inom kontrollprogrammet och riktvärdena i prövotidsvillkoret. Dessutom utvärderades effektiviteten i varje enskilt reningssteg, den kemiska reningen och kol- och torvfiltret. För optimering av den kemiska reningen testades lakvattnet först på lab med olika fällnings- och flockningsmedel. Uppgiften var att öka partikelfällningen och reducera metallerna arsenik, kadmium, krom, kvicksilver, bly, koppar, nickel och zink i lakvattnet genom svepkoagulering och hydroxidfällning.</p><p>Utifrån resultaten på lab testades sedan fällningen i full skala, pH höjdes i flockningsbassängen och så även dosen fällningsmedel till vattnet. En inledande undersökning av regleringen av lutdosering gjordes med några stegsvarsexperiment.</p><p>Lakvatten innehöll låga halter av organiskt material och närsalterna kväve och fosfor, medan de halogena ämnena (AOX) och tungmetallerna nickel och koppar var höga. De båda reningsstegen, kemisk rening och kol- och torvfiltret var generellt bra på att rena föroreningar i höga koncentrationer men var sämre vid låga. Kol- och torvfiltret ökade koncentrationen i vattnet av arsenik genom materialets interna läckage. I fällningsförsöken gav järnkloriden, PlusJÄRN och den anjoniska polyakrylamiden, Fennopol A bäst resultat i att avskilja metaller. Järnsulfaten PurFect gav näst bäst resultat och valdes istället för järnkloriden för vidare försök då kloridhalten i lakvattnet redan var högt. Optimalt pH för arsenik, zink, koppar och nickel var inom pHintervallet 9 och 9,5. Fällningen i full skala ute i verket visade ett bättre resultat vid tillsatt lut och ett högre pH (pH 9) i flockningsbassängen. Däremot betydde en höjning av fällningskemikalien från 202 mg/l PurFect till 225 mg/l enbart en ökad kemikaliekostnad och en överdosering. Det visade sig dessutom att lutregleringen var svängig och på gränsen till instabil. Regulatorn bör därför ses över så att risken för ökad kemikaliekonsumption och kostnader minskar.</p>
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Zobrazení šachů pomocí sledování paprsku / Rendering Chess Using Ray TracingTurek, Vojtěch Unknown Date (has links)
This work deals with ray tracing technique, focused to rendering of chess scene with procedural texture materials. Based on this knowledge, program that displays specified chess set is designed.
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Membrane distillation with porous metal hollow fibers for the concentration of thermo-sensitive solutions / Distillation membranaire avec des fibres creuses métalliques pour la concentration des solutions thermo-sensiblesShukla, Sushumna 18 December 2014 (has links)
Cette thèse présente une approche originale du procédé de distillation membranaire avec balayage gazeux pour la concentration des solutions thermosensibles (SGMD). Pour ce faire, un nouveau contacteur membranaire avec des fibres creuses métalliques a été conçu afin réaliser le procédé de distillation à basse température. La chaleur nécessaire au procédé est produite au niveau des fibres par effet Joule, plutôt qu'à partir de chaleur latente de la phase aqueuse. La génération localisée de la chaleur a comme conséquence une réduction du phénomène de polarisation de la température. Des fibres creuses en acier inoxydable ont été synthétisées avec les propriétés structurales appropriées et une bonne résistance mécanique. La surface des pores des fibres a été rendue hydrophobe par le dépôt d'une fine couche d'un élastomère. En outre, une nouvelle méthode « verte » a été développée pour fabriquer des fibres creuses en alumine et acier inoxydable. Cette méthode est basée sur la gélification ionique des bio-polymères et ne n'utilise pas des solvants nocifs. L'étude expérimentale détaillée du SGMD a permis de déterminer l'influence de différents paramètres opérationnels sur les performances du procédé. Il a été démontré que l'effet Joule permet d'améliorer le flux et l'efficacité de la séparation non seulement pour le SGMD mais aussi pour la pervaporation. / This thesis presents an original approach for the concentration of thermo-sensitive solutions: the Sweep Gas Membrane Distillation (SGMD) process. A new membrane contactor with metallic hollow fibers has been designed and allows the distillation process to be operational at low temperature. Heat is generated in the fibers by the Joule effect, rather than being supplied as latent heat in the liquid bulk. The localized generation of heat results in a reduction of temperature polarization phenomena. The stainless-steel hollow fiber membranes have been synthetized with appropriate structural properties and sufficient mechanical strength. The pore surface of the fibers has been made hydrophobic by the deposition of a thin layer of an elastomer. Moreover, a novel and green method is presented to fabricate alumina and stainless-steel hollow fibers. This method is based on ionic gelation of a biopolymer and completely avoids the use of harmful solvents. By a detailed experimental study of the SGMD the influence of different operational parameters on the process performance has been investigated. The improvements in the flux and the separation efficiency using Joule effect have been successfully demonstrated, even in the case of pervaporation.
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Passage à l'échelle pour les contraintes d'ordonnancement multi-ressources / Scalable multi-dimensional resources scheduling constraintsLetort, Arnaud 28 October 2013 (has links)
La programmation par contraintes est une approche régulièrement utilisée pour résoudre des problèmes combinatoires d’origines diverses. Dans cette thèse nous nous focalisons sur les problèmes d’ordonnancement cumulatif. Un problème d’ordonnancement consiste à déterminer les dates de débuts et de fins d’un ensemble de tâches, tout en respectant certaines contraintes de capacité et de précédence. Les contraintes de capacité concernent aussi bien des contraintes cumulatives classiques où l’on restreint la somme des hauteurs des tâches intersectant un instant donné, que des contraintes cumulatives colorées où l’on restreint le nombre maximum de couleurs distinctes prises par les tâches. Un des objectifs récemment identifiés pour la programmation par contraintes est de traiter des problèmes de grandes tailles, habituellement résolus à l’aide d’algorithmes dédiés et de métaheuristiques. Par exemple, l’utilisation croissante de centres de données virtualisés laisse apparaitre des problèmes d’ordonnancement et de placement multi-dimensionnels de plusieurs milliers de tâches. Pour atteindre cet objectif, nous utilisons l’idée de balayage synchronisé considérant simultanément une conjonction de contraintes cumulative et des précédences, ce qui nous permet d’accélérer la convergence au point fixe. De plus, de ces algorithmes de filtrage nous dérivons des procédures gloutonnes qui peuvent être appelées à chaque nœud de l’arbre de recherche pour tenter de trouver plus rapidement une solution au problème. Cette approche permet de traiter des problèmes impliquant plus d’un million de tâches et 64 ressources cumulatives. Ces algorithmes ont été implémentés dans les solveurs de contraintes Choco et SICStus, et évalués sur divers problèmes déplacement et d’ordonnancement. / Constraint programming is an approach often used to solve combinatorial problems in different application areas. In this thesis we focus on the cumulative scheduling problems. A scheduling problem is to determine the starting dates of a set of tasks while respecting capacity and precedence constraints. Capacity constraints affect both conventional cumulative constraints where the sum of the heights of tasks intersecting a given time point is limited, and colored cumulative constraints where the number of distinct colors assigned to the tasks intersecting a given time point is limited. A newly identified challenge for constraint programming is to deal with large problems, usually solved by dedicated algorithms and metaheuristics. For example, the increasing use of virtualized datacenters leads to multi dimensional placement problems of thousand of jobs. Scalability is achieved by using a synchronized sweep algorithm over the different cumulative and precedence constraints that allows to speed up convergence to the fix point. In addition, from these filtering algorithms we derive greedy procedures that can be called at each node of the search tree to find a solution more quickly. This approach allows to deal with scheduling problems involving more than one million jobs and 64 cumulative resources. These algorithms have been implemented within Choco and SICStussolvers and evaluated on a variety of placement and scheduling problems.
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Optimisation d'une source vibratoire pour la détection des cavités souterraines par sismique réflexion haute résolution / Optimization of a vibratory source for cavity detection by high resolution seismicKosecki, Arkadiusz 07 December 2009 (has links)
L’objectif principal de cette thèse est de développer et d’optimiser les outils d’acquisition de la technique de Sismique réflexion haute résolution (SHR) afin d’améliorer ses performances pour la détection des cavités souterraines. Il est communément admis que l’imagerie SHR est d’autant plus complexe que la profondeur de la cible est petite. Les travaux menés dans le cadre de cette thèse devraient remédier à certains problèmes les plus critiques identifiés lors des applications de la SHR.L’utilisation des sources vibratoires présente des avantages indéniables (non destructivité, contrôle du signal émis...) par rapport aux sources « classiques » (i.e. impulsionnelles, destructives) mais leur application optimale nécessite un choix correct du signal émis.Ainsi, les travaux de recherche réalisés ont permis de (1) développer un système complet de pilotage par ordinateur d’une mini-source vibratoire destinée à l’imagerie SHR, (2) développer une méthode de génération de signaux émis. En établissant un lien entre le signal d’entrée et l’image sismique obtenue, cette procédure offre la possibilité à l’utilisateur de choisir le signal émis en fonction des conditions de terrain, et des objectifs des mesures, (3) tester le fonctionnement du système développé avec plusieurs mini-vibrateurs.Le système développé ainsi est testé et validé dans les tests à petite échelle. Ensuite, il a été utilisé dans les conditions réelles avec l’objectif « détection des cavités » dans le contexte salin (anciennes mines de sel en Lorraine, profondeur : 160 m - 180 m) et les marnières de Haute Normandie (anciennes carrières de craie, profondeur : 15 m - 45 m) / The main objective of this thesis is to develop and optimize the acquisition tools for High Resolution Reflection Seismic (HRS) technique in order to improve its performances in the detection of underground cavities. It is commonly admitted that HRS imaging becomes more complicated with when the depth of interest is decreased. The work carried out in the frame of this thesis aims to bring solutions to some of the most critical problems identified in application of the HRS.The vibratory sources show undeniable advantages (non destructivity, controllable output signal) over “classic” (impulsive, destructive) sources. However, the optimal use of these sources depends on the proper choice of emitted signal.Thus, the research work carried out resulted in (1) development of a complete, computer-based vibrator control system allowing piloting small vibratory source intended to use for HRS surveys, (2) development of a method for generating the source signal. The proposed procedure links the entry signal with seismic image and thus allows the choice of the signal in terrain conditions and with regard to the measurement goals, (3) extensive testing of the developed system with several portable vibratory sources.The developed system was tested and validated in small-scale tests. Afterwards it was used in real conditions with the goal of “cavity detection” in salt-mining context (old salt mines in Lorraine region at depths between 160 m and 180 m) as well as in chalk-mining area (ancient marl-pit quarries in the Normandy region at depths 15 m - 45 m)
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Tracing selection and adaptation along an environmental gradient in Populus tremulaHall, David January 2009 (has links)
The distribution of the expressed genotype is moved around in the population over time byevolution. Natural selection is one of the forces that act on the phenotype to change the patterns ofnucleotide variation underlying those distributions. How the phenotype changes over aheterogeneous environment describes the type of evolutionary force acting on this trait and thisshould be reflected in the variation at loci underlying this trait. While the variation in phenotypesand at the nucleotide level in a population indicates the same evolutionary force, it does notnecessarily mean that they are connected. In natural populations the continuous shifting of geneticmaterial through recombination events break down possible associations between loci facilitates theexamination of possible causal loci to single base pair differences in DNA-sequences. Connecting thegenotype and the phenotype thus provides an important step in the understanding the geneticarchitecture of complex traits and the forces that shape the observed patterns.This thesis examines the European aspen, Populus tremula, sampled from subpopulations overan extensive latitudinal gradient covering most of Sweden. Results show a clear geneticdifferentiation in the timing of bud set, a measure of the autumnal cessation of growth, betweendifferent parts of Sweden pointing at local adaptation. In the search for candidate genes thatunderlie the local adaptation found, most genes (25) in the photoperiodic gene network wereexamined for signals of selection. Genes in the photoperiodic network show an increase in theheterogeneity of differentiation between sampled subpopulations in Sweden. Almost half (12) of theexamined genes are under some form of selection. Eight of these genes show positive directionalselection on protein evolution and the gene that code for a photoreceptor, responsible for mediatingchanging light conditions to downstream targets in the network, has the hallmarks of a selectivesweep. The negative correlation between positive directional selection and synonymous diversityindicates that the majority of the photoperiod gene network has undergone recurrent selectivesweeps. A phenomenon that likely has occurred when P. tremula has readapted to the northern lightregimes during population expansion following retracting ice between periods of glaciations. Two ofthe genes under selection also have single nucleotide polymorphisms (SNP) that associate with budset, two in the PHYB2 gene and one in the LHY2 gene. Furthermore, there is an additional SNP inLHY1 that explain part of the variation in timing of bud set, despite the lack of a signal of selection atthe LHY1 gene. Together these SNPs explain 10-15% of the variation in the timing of bud set and 20-30% more if accounting for the positive co-variances between SNPs. There is thus rather extensiveevidence that genes in the photoperiod gene network control the timing of bud set, and reflect localadaptation in this trait.
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Optimering av den kemiska reningen vid Fläskebo deponi / Optimization of the Chemical Treatment at Fläskebo LandfillNilsson, Anna January 2006 (has links)
Landfill leachate contains a variety of contaminants and is created when rainwater percolates the landfill. For landfill management the leachate is the main issue that can cause problems to the environment. At the landfill of Fläskebo, Renova AB treats the leachate in a local treatment plant. The treatment consists of a chemical treatment step with chemical precipitation, flocculation, sedimentation and filtration, and a final step with a carbon and peat filter. Renova has to ensure that the condition of the leachate reaches the regulation set for the landfill before it is released to the recipient. This regulation has not yet been established and a final suggestion will be given to the county administrative board in spring 2006. In this master thesis the chemical treatment of Fläskebo is optimized. A comparison between the control program and the regulation was made to estimate the contamination of the leachate. Also the effectiveness of the two steps is evaluated. For optimization, the leachate was first tested in a laboratory with different coagulants and flocculants. The purpose was to increase the precipitation of particles and metals; arsenic, cadmium, chromium, mercury, lead, copper, nickel and zinc through sweep floc coagulation and hydroxide precipitation. After the laboratory tests the precipitation was tested in the treatment plant with higher pH and coagulant. Also the process control for sodium hydroxide was examined. The leachate had a small content of organic matter and nutrients, but had a large content of halogenated substances (AOX) and the heavy metals nickel and copper. High concentrations of contaminants were reduced better than low concentrations in the two treatment steps. The carbon and peat filter material also caused an increase of the arsenic content in the leachate after filtration. From the laboratory work the results showed a better reduction of metals with iron-chloride, PlusJÄRN and the anjonic polyacrylamid, Fennopol A. Because of the high content of chloride the iron- sulphate, PurFect was chosen for further tests. The optimal pH for the heavy metals arsenic, zinc, copper and nickel was between pH 9 and 9, 5. The precipitation in the treatment plant showed better results with sodium hydroxide and a higher pH, pH 9 in the flocculation basin. An increase of the coagulant PurFect from 202 mg/l to 225 mg/l meant only a higher chemical cost. The process control of sodium hydroxide showed an oscillating and unstable control performance, which may lead to a higher consumption of chemicals. / Lakvatten har varierande föroreningsgrad och karaktär och bildas då regnvatten perkolerar igenom en deponi. För en deponiverksamhet är lakvattnet den huvudsakliga påverkan på den omgivande miljön. Vid en av Renova ABs deponier, Fläskebo, utanför Göteborg sker reningen av vattnet i den lokala reningsanläggningen. Reningen består av kemisk fällning, flockning, sedimentering och filtrering samt ett kol- och torvfilter. För att få släppa ut lakvattnet har Renova just nu ett prövotidsvillkor på lakvattnets kvalité och till våren 2006 skall förslag på slutgiltiga utsläppsvillkor lämnas till Länsstyrelsen. I detta examensarbete har den kemiska reningen av lakvattnet från Fläskebo optimerats utifrån lakvattnets karaktär. Lakvattnets föroreningsgrad bedömdes efter en jämförelse av analysresultatet inom kontrollprogrammet och riktvärdena i prövotidsvillkoret. Dessutom utvärderades effektiviteten i varje enskilt reningssteg, den kemiska reningen och kol- och torvfiltret. För optimering av den kemiska reningen testades lakvattnet först på lab med olika fällnings- och flockningsmedel. Uppgiften var att öka partikelfällningen och reducera metallerna arsenik, kadmium, krom, kvicksilver, bly, koppar, nickel och zink i lakvattnet genom svepkoagulering och hydroxidfällning. Utifrån resultaten på lab testades sedan fällningen i full skala, pH höjdes i flockningsbassängen och så även dosen fällningsmedel till vattnet. En inledande undersökning av regleringen av lutdosering gjordes med några stegsvarsexperiment. Lakvatten innehöll låga halter av organiskt material och närsalterna kväve och fosfor, medan de halogena ämnena (AOX) och tungmetallerna nickel och koppar var höga. De båda reningsstegen, kemisk rening och kol- och torvfiltret var generellt bra på att rena föroreningar i höga koncentrationer men var sämre vid låga. Kol- och torvfiltret ökade koncentrationen i vattnet av arsenik genom materialets interna läckage. I fällningsförsöken gav järnkloriden, PlusJÄRN och den anjoniska polyakrylamiden, Fennopol A bäst resultat i att avskilja metaller. Järnsulfaten PurFect gav näst bäst resultat och valdes istället för järnkloriden för vidare försök då kloridhalten i lakvattnet redan var högt. Optimalt pH för arsenik, zink, koppar och nickel var inom pHintervallet 9 och 9,5. Fällningen i full skala ute i verket visade ett bättre resultat vid tillsatt lut och ett högre pH (pH 9) i flockningsbassängen. Däremot betydde en höjning av fällningskemikalien från 202 mg/l PurFect till 225 mg/l enbart en ökad kemikaliekostnad och en överdosering. Det visade sig dessutom att lutregleringen var svängig och på gränsen till instabil. Regulatorn bör därför ses över så att risken för ökad kemikaliekonsumption och kostnader minskar.
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Range Resolution Improvement Of Fmcw RadarsKurt, Sinan 01 September 2007 (has links) (PDF)
Frequency Modulated Continuous Wave (FMCW) radar has wide application areas in both civil and military use. The range resolution is a critical concept for these FMCW radars as for the other radar types. There are theoretical restrictions in the range resolution. In addition, the non-ideal properties of the modules used in the systems negatively affects the range resolution. The transmitter leakage, non-linear frequency sweep, FM to AM distortion and measurement errors are some of the critical non-ideal properties. The problems arising from these non-ideal properties further restrict the range resolution of FMCW radars. Another important concept for the range resolution that can be obtained from FMCW radars is the signal processing method. This thesis deals with the non-ideal properties of the system modules and techniques to reduce their effects on the range resolution. Furthermore, the signal processing methods used for FMCW radar signals and the possible improvement techniques for these methods are discussed. Moreover, a simple signal processing unit called zero crossing counter which can be used for short range FMCW radars is implemented and range resolution performance of this zero crossing counter is investigated by carrying out measurements on a prototype FMCW radar at 2200MHz.
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Genetic consequences of directional selection in <em>Arabidopsis lyrata</em>Toivainen, T. (Tuomas) 01 December 2014 (has links)
Abstract
Plants and animals colonized Northern Europe after the last Ice Age from different refugia, not covered by the ice sheet. Many plants, such as the northern rock cress (Arabidopsis lyrata ssp. petraea) adapted to the short growing season in the North. We thus expect that colonization of the new environment was accompanied by directional selection for traits conferring this adaptation. In this thesis I studied whether recent directional selection can be detected in two important genes, PHYTOCHROME A (PHYA) and FLOWERING LOCUS C1 (FLC1), related to the flowering time pathway. To detect directional selection, I compared DNA sequence variation from the samples of a southern (Plech, Germany) and a northern (Spiterstulen, Norway) population. I also studied the current response potential to changing conditions in the marginal Spiterstulen population. Adaptation potential was characterized by assessing plasticity and amount of additive genetic variation, focusing on flowering traits. In addition, associations of 21 flowering time candidate genes for phenological and fitness traits were studied.
There were several lines of evidence for recent directional selection in both candidate genes, PHYA and FLC1, in the northern Spiterstulen population Variation was strongly reduced around both genes and in addition they were highly differentiated between populations. In the Spiterstulen population there was a remarkable reduction in additive genetic variation for flowering traits, for instance when compared with morphological traits. On the other hand, phenological traits showed high plasticity. Some of the photoperiodic pathway genes showed association to flowering or reproductive fitness.
The results suggest that directional selection during the colonization of the northern areas has impacted the two studied genes. Genetic changes were likely involved in altered photoperiodic and vernalization responses which might be adaptive for a short growing season. Further, directional selection was probably responsible for reducing additive genetic variation in flowering traits. Because there was only little genetic variation, adaptation to future environmental change of the marginal Spiterstulen population is likely to rely largely on plastic reactions to environmental signals, or tracking the environment by dispersal. / Tiivistelmä
Kasvit ja eläimet levittäytyivät Pohjois-Eurooppaan viimeisen jääkauden jälkeen mannerjäätikön ulkopuolella jääneistä refugioista. Useat kasvit, kuten idänpitkäpalko (Arabidopsis lyrata ssp. petraea) sopeutuivat pohjoisen lyhyeen kasvukauteen. On syytä olettaa, että suuntaava valinta vaikutti sopeutumisessa tärkeisiin ominaisuuksiin. Tässä väitöskirjassa tutkin voidaanko suuntaavan valinnan aiheuttamia jalanjälkiä löytää kahdesta tärkeästä kukkimisaikageenistä, FYTOKROMI A (PHYA) ja FLOWERING LOCUS C1 (FLC1) geeneistä. Tätä varten vertasin DNA sekvenssimuuntelua pohjoisessa (Norja) ja eteläisessä (Saksa) populaatiossa, kiinnittäen erityisesti huomiota geneettisen muuntelun määrään ja erilaistumiseen. Lisäksi tutkin miten Spiterstulenin reunapopulaatio voi vastata tulevaisuudessa muuttuvaan ympäristöön. Sopeutumispotentiaalia arvioitiin sekä fenotyyppisen plastisuuden että additiivisen geneettisen muuntelun määrällä. Lisäksi tutkin miten vaihtelu 21 kukkimisaikageenissä liittyy fenologisiin ja kelpoisuusominaisuuksiin.
Useat merkit viittasivat siihen, että suuntaava valinta oli vaikuttanut kummassakin tutkitussa geenissä. Muuntelu oli vähentynyt voimakkaasti kumpaakin geeniä ympäröiviltä kromosomialueilta, jotka olivat myös selkeästi erilaistuneet. Additiivinen geneettinen muuntelu oli selvästi vähentynyt kukkimisominaisuuksissa verrattuna morfologisiin ominaisuuksiin, mahdollisesti suuntaavan valinnan johdosta. Kukkimisominaisuudet olivat kuitenkin plastisia. Jotkin valojaksoreitin geenit vaikuttivat sekä kukkimiseen että lisääntymiskykyyn.
Nämä tulokset osoittavat että suuntaava valinta vaikutti kahteen tutkittuun geeniin pohjoiseen levittäytymisen aikana. Geneettiset muutokset liittyivät todennäköisesti muuttuneisiin valojakso-, ja vernalisaatiovasteisiin, jotka saattoivat edistää sopeutumista lyhyeen kasvukauteen. Koska geneettistä muuntelua oli vain hyvin vähän, fenotyyppisellä plastisuudella on todennäköisesti tärkeä rooli sopeutumisessa muuttuvaan ympäristöön Spiterstulenin reunapopulaatiossa.
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Post-glacial colonization, demographic history, and selection in <em>Arabidopsis lyrata</em>:genome-wide and candidate gene based approachMattila, T. (Tiina) 31 October 2017 (has links)
Abstract
Demographic history and natural selection are central forces shaping the genetic diversity of populations. Knowledge on these forces increases understanding of processes shaping genetic variability of populations. In this PhD thesis I investigated demographic history and selection in multiple populations of Arabidopsis lyrata, an outcrossing herbaceous plant species of the Brassicaceae family. Due to its wide distribution in the temperate and boreal regions, A. lyrata serves as a good model system to study population genetic consequences of colonization of northern latitudes. The first aim of this study was to characterize the demographic and colonization history of the species using site frequency spectra estimated from whole-genome diversity data. Another aim was to detect genetic loci targeted by recent selective sweeps at genome-wide scale as well as at candidate flowering time genes. Patterns of genome-wide selection at linked sites (linked selection) were also compared between populations of Capsella grandiflora and A. lyrata with contrasting demographic histories.
Evidence for strong effective population size decline in the past few hundred thousand years was detected in A. lyrata populations species-wide. This study also suggests recent Scandinavian colonization from an unknown refugium, distinct from the Central European source population. Selection analyses revealed loci targeted by positive selection in two Scandinavian lineages after the recent population split as well as selective sweeps in flowering time genes in the colonizing populations. In comparison with the studied C. grandiflora population, the Norwegian A. lyrata population had weaker purifying selection and no evidence for reduction of diversity around genes was found. This thesis offers novel information on species colonization history and its genome-wide effects, which is important for understanding the framework of local adaptation. / Tiivistelmä
Populaation demografinen historia ja luonnonvalinta ovat keskeisiä populaation perinnöllisen muuntelun muokkaajia. Näiden tekijöiden tutkimus on tärkeää eliöiden sopeutumisen ymmärtämiselle. Tässä väitöskirjassa tutkin demografista historiaa ja valintaa monivuotisen ristisiittoiseen ruohovartisen Brassicaceae-heimon kasvilajin idänpitkäpalon (Arabidopsis lyrata) useissa eri populaatioissa. Idänpitkäpalko on erinomainen mallilaji pohjoiseen ympäristöön sopeutumisen tutkimukseen, koska sen toisistaan eristäytyneet paikalliset populaatiot ovat levittäytyneet laajalle boreaalisella ja lauhkealla ilmastovyöhykkeellä. Tutkimuksen tarkoituksena oli luonnehtia populaatioiden demografista historiaa ja kolonisaatioreittejä käyttäen koko perimän laajuisesta muunteluaineistosta estimoituja alleelifrekvenssispektrejä. Lisäksi koko perimän laajuista aineistoa sekä kukkimisaikaa ohjaavien geenien sekvenssejä käytettiin positiivisen luonnonvalinnan merkkien tunnistukseen. Genominlaajuista kytkeytynyttä valintaa vertailtiin toiseen ristisiittoiseen Brassicaceae-heimon lajin Capsella grandifloran populaatioon, jonka demografinen historia poikkeaa huomattavasti tutkituista idänpitkäpalon populaatioista.
Tutkimuksessa havaittiin, että kaikissa tutkituissa idänpitkäpalon populaatioissa tehollinen populaatiokoko oli pienentynyt viimeisen muutaman sadantuhannen vuoden aikana. Kolonisaatiohistorian tarkastelu osoitti, että idänpitkäpalon skandinaaviset populaatiot ovat todennäköisesti peräisin keskieurooppalaisesta refugiosta erillisestä läntisestä refugiosta. Skandinavian kolonisaation yhteydessä vaikuttaneen positiivisen luonnonvalinnan merkkejä havaittiin useissa eri genomin osissa sekä erityisesti valojaksoa mittaavissa geeneissä. Tämä kertoo erilaisiin valojaksoihin sopeutumisen tärkeydestä skandinaavisen kolonisaation yhteydessä. Verrattuna tutkittuun C. grandifloran populaatioon, idänpitkäpalolla puhdistavan valinnan havaittiin olevan heikompaa ja muuntelun vähenemistä geenien ympärillä ei havaittu. Tämä tutkimus tarjoaa uutta tietoa Skandinavian kolonisaatiohistoriasta ja sen genominlaajuisista vaikutuksista. Tutkimuksessa tuotettua tietoa voidaan hyödyntää paikallisen sopeutumisen ymmärtämisessä.
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