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Intracellular photoactivation of caged cGMP induces myosin II and actin responses in motile cells
(2013)
Cyclic GMP (cGMP) is a ubiquitous second messenger in eukaryotic cells. It is assumed to regulate the association of myosin II with the cytoskeleton of motile cells. When cells of the social amoeba Dictyostelium discoideum are exposed to chemoattractants or to increased osmotic stress, intracellular cGMP levels rise, preceding the accumulation of myosin II in the cell cortex. To directly investigate the impact of intracellular cGMP on cytoskeletal dynamics in a living cell, we released cGMP inside the cell by laser-induced photo-cleavage of a caged precursor. With this approach, we could directly show in a live cell experiment that an increase in intracellular cGMP indeed induces myosin II to accumulate in the cortex. Unexpectedly, we observed for the first time that also the amount of filamentous actin in the cell cortex increases upon a rise in the cGMP concentration, independently of cAMP receptor activation and signaling. We discuss our results in the light of recent work on the cGMP signaling pathway and suggest possible links between cGMP signaling and the actin system.
Galaxy clusters are the largest known gravitationally bound objects, their study is important for both an intrinsic understanding of their systems and an investigation of the large scale structure of the universe. The multi- component nature of galaxy clusters offers multiple observable signals across the electromagnetic spectrum. At X-ray wavelengths, galaxy clusters are simply identified as X-ray luminous, spatially extended, and extragalactic sources. X-ray observations offer the most powerful technique for constructing cluster catalogues. The main advantages of the X-ray cluster surveys are their excellent purity and completeness and the X-ray observables are tightly correlated with mass, which is indeed the most fundamental parameter of clusters. In my thesis I have conducted the 2XMMi/SDSS galaxy cluster survey, which is a serendipitous search for galaxy clusters based on the X-ray extended sources in the XMM-Newton Serendipitous Source Catalogue (2XMMi-DR3). The main aims of the survey are to identify new X-ray galaxy clusters, investigate their X-ray scaling relations, identify distant cluster candidates, and study the correlation of the X-ray and optical properties. The survey is constrained to those extended sources that are in the footprint of the Sloan Digital Sky Survey (SDSS) in order to be able to identify the optical counterparts as well as to measure their redshifts that are mandatory to measure their physical properties. The overlap area be- tween the XMM-Newton fields and the SDSS-DR7 imaging, the latest SDSS data release at the starting of the survey, is 210 deg^2. The survey comprises 1180 X-ray cluster candidates with at least 80 background-subtracted photon counts, which passed the quality control process. To measure the optical redshifts of the X-ray cluster candidates, I used three procedures; (i) cross-matching these candidates with the recent and largest optically selected cluster catalogues in the literature, which yielded the photometric redshifts of about a quarter of the X-ray cluster candidates. (ii) I developed a finding algorithm to search for overdensities of galaxies at the positions of the X-ray cluster candidates in the photometric redshift space and to measure their redshifts from the SDSS-DR8 data, which provided the photometric redshifts of 530 groups/clusters. (iii) I developed an algorithm to identify the cluster candidates associated with spectroscopically targeted Luminous Red Galaxies (LRGs) in the SDSS-DR9 and to measure the cluster spectroscopic redshift, which provided 324 groups and clusters with spectroscopic confirmation based on spectroscopic redshift of at least one LRG. In total, the optically confirmed cluster sample comprises 574 groups and clusters with redshifts (0.03 ≤ z ≤ 0.77), which is the largest X-ray selected cluster catalogue to date based on observations from the current X-ray observatories (XMM-Newton, Chandra, Suzaku, and Swift/XRT). Among the cluster sample, about 75 percent are newly X-ray discovered groups/clusters and 40 percent are new systems to the literature. To determine the X-ray properties of the optically confirmed cluster sample, I reduced and analysed their X-ray data in an automated way following the standard pipelines of processing the XMM-Newton data. In this analysis, I extracted the cluster spectra from EPIC(PN, MOS1, MOS2) images within an optimal aperture chosen to maximise the signal-to-noise ratio. The spectral fitting procedure provided the X-ray temperatures kT (0.5 - 7.5 keV) for 345 systems that have good quality X-ray data. For all the optically confirmed cluster sample, I measured the physical properties L500 (0.5 x 10^42 – 1.2 x 10^45 erg s-1 ) and M500 (1.1 x 10^13 – 4.9 x 10^14 M⊙) from an iterative procedure using published scaling relations. The present X-ray detected groups and clusters are in the low and intermediate luminosity regimes apart from few luminous systems, thanks to the XMM-Newton sensitivity and the available XMM-Newton deep fields The optically confirmed cluster sample with measurements of redshift and X-ray properties can be used for various astrophysical applications. As a first application, I investigated the LX - T relation for the first time based on a large cluster sample of 345 systems with X-ray spectroscopic parameters drawn from a single survey. The current sample includes groups and clusters with wide ranges of redshifts, temperatures, and luminosities. The slope of the relation is consistent with the published ones of nearby clusters with higher temperatures and luminosities. The derived relation is still much steeper than that predicted by self-similar evolution. I also investigated the evolution of the slope and the scatter of the LX - T relation with the cluster redshift. After excluding the low luminosity groups, I found no significant changes of the slope and the intrinsic scatter of the relation with redshift when dividing the sample into three redshift bins. When including the low luminosity groups in the low redshift subsample, I found its LX - T relation becomes after than the relation of the intermediate and high redshift subsamples. As a second application of the optically confirmed cluster sample from our ongoing survey, I investigated the correlation between the cluster X-ray and the optical parameters that have been determined in a homogenous way. Firstly, I investigated the correlations between the BCG properties (absolute magnitude and optical luminosity) and the cluster global proper- ties (redshift and mass). Secondly, I computed the richness and the optical luminosity within R500 of a nearby subsample (z ≤ 0.42, with a complete membership detection from the SDSS data) with measured X-ray temperatures from our survey. The relation between the estimated optical luminosity and richness is also presented. Finally, the correlation between the cluster optical properties (richness and luminosity) and the cluster global properties (X-ray luminosity, temperature, mass) are investigated.
LCST-type synthetic thermoresponsive polymers can reversibly respond to certain stimuli in aqueous media with a massive change of their physical state. When fluorophores, that are sensitive to such changes, are incorporated into the polymeric structure, the response can be translated into a fluorescence signal. Based on this idea, this thesis presents sensing schemes which transduce the stimuli-induced variations in the solubility of polymer chains with covalently-bound fluorophores into a well-detectable fluorescence output. Benefiting from the principles of different photophysical phenomena, i.e. of fluorescence resonance energy transfer and solvatochromism, such fluorescent copolymers enabled monitoring of stimuli such as the solution temperature and ionic strength, but also of association/disassociation mechanisms with other macromolecules or of biochemical binding events through remarkable changes in their fluorescence properties. For instance, an aqueous ratiometric dual sensor for temperature and salts was developed, relying on the delicate supramolecular assembly of a thermoresponsive copolymer with a thiophene-based conjugated polyelectrolyte. Alternatively, by taking advantage of the sensitivity of solvatochromic fluorophores, an increase in solution temperature or the presence of analytes was signaled as an enhancement of the fluorescence intensity. A simultaneous use of the sensitivity of chains towards the temperature and a specific antibody allowed monitoring of more complex phenomena such as competitive binding of analytes. The use of different thermoresponsive polymers, namely poly(N-isopropylacrylamide) and poly(meth)acrylates bearing oligo(ethylene glycol) side chains, revealed that the responsive polymers differed widely in their ability to perform a particular sensing function. In order to address questions regarding the impact of the chemical structure of the host polymer on the sensing performance, the macromolecular assembly behavior below and above the phase transition temperature was evaluated by a combination of fluorescence and light scattering methods. It was found that although the temperature-triggered changes in the macroscopic absorption characteristics were similar for these polymers, properties such as the degree of hydration or the extent of interchain aggregations differed substantially. Therefore, in addition to the demonstration of strategies for fluorescence-based sensing with thermoresponsive polymers, this work highlights the role of the chemical structure of the two popular thermoresponsive polymers on the fluorescence response. The results are fundamentally important for the rational choice of polymeric materials for a specific sensing strategy.
Passive plant actuators have fascinated many researchers in the field of botany and structural biology since at least one century. Up to date, the most investigated tissue types in plant and artificial passive actuators are fibre-reinforced composites (and multilayered assemblies thereof) where stiff, almost inextensible cellulose microfibrils direct the otherwise isotropic swelling of a matrix. In addition, Nature provides examples of actuating systems based on lignified, low-swelling, cellular solids enclosing a high-swelling cellulosic phase. This is the case of the Delosperma nakurense seed capsule, in which a specialized tissue promotes the reversible opening of the capsule upon wetting. This tissue has a diamond-shaped honeycomb microstructure characterized by high geometrical anisotropy: when the cellulosic phase swells inside this constraining structure, the tissue deforms up to four times in one principal direction while maintaining its original dimension in the other. Inspired by the example of the Delosoperma nakurense, in this thesis we analyze the role of architecture of 2D cellular solids as models for natural hygromorphs. To start off, we consider a simple fluid pressure acting in the cells and try to assess the influence of several architectural parameters onto their mechanical actuation. Since internal pressurization is a configurational type of load (that is the load direction is not fixed but it “follows” the structure as it deforms) it will result in the cellular structure acquiring a “spontaneous” shape. This shape is independent of the load but just depends on the architectural characteristics of the cells making up the structure itself. Whereas regular convex tiled cellular solids (such as hexagonal, triangular or square lattices) deform isotropically upon pressurization, we show through finite element simulations that by introducing anisotropic and non-convex, reentrant tiling large expansions can be achieved in each individual cell. The influence of geometrical anisotropy onto the expansion behaviour of a diamond shaped honeycomb is assessed by FEM calculations and a Born lattice approximation. We found that anisotropic expansions (eigenstrains) comparable to those observed in the keels tissue of the Delosoperma nakurense are possible. In particular these depend on the relative contributions of bending and stretching of the beams building up the honeycomb. Moreover, by varying the walls’ Young modulus E and internal pressure p we found that both the eigenstrains and 2D elastic moduli scale with the ratio p/E. Therefore the potential of these pressurized structures as soft actuators is outlined. This approach was extended by considering several 2D cellular solids based on two types of non-convex cells. Each honeycomb is build as a lattice made of only one non-convex cell. Compared to usual honeycombs, these lattices have kinked walls between neighbouring cells which offers a hidden length scale allowing large directed deformations. By comparing the area expansion in all lattices, we were able to show that less convex cells are prone to achieve larger area expansions, but the direction in which the material expands is variable and depends on the local cell’s connectivity. This has repercussions both at the macroscopic (lattice level) and microscopic (cells level) scales. At the macroscopic scale, these non-convex lattices can experience large anisotropic (similarly to the diamond shaped honeycomb) or perfectly isotropic principal expansions, large shearing deformations or a mixed behaviour. Moreover, lattices that at the macroscopic scale expand similarly can show quite different microscopic deformation patterns that include zig-zag motions and radical changes of the initial cell shape. Depending on the lattice architecture, the microscopic deformations of the individual cells can be equal or not, so that they can build up or mutually compensate and hence give rise to the aforementioned variety of macroscopic behaviours. Interestingly, simple geometrical arguments involving the undeformed cell shape and its local connectivity enable to predict the results of the FE simulations. Motivated by the results of the simulations, we also created experimental 3D printed models of such actuating structures. When swollen, the models undergo substantial deformation with deformation patterns qualitatively following those predicted by the simulations. This work highlights how the internal architecture of a swellable cellular solid can lead to complex shape changes which may be useful in the fields of soft robotics or morphing structures.
Numerical simulations of galaxy formation and observational Galactic Astronomy are two fields of research that study the same objects from different perspectives. Simulations try to understand galaxies like our Milky Way from an evolutionary point of view while observers try to disentangle the current structure and the building blocks of our Galaxy. Due to great advances in computational power as well as in massive stellar surveys we are now able to compare resolved stellar populations in simulations and in observations. In this thesis we use a number of approaches to relate the results of the two fields to each other. The major observational data set we refer to for this work comes from the Radial Velocity Experiment (RAVE), a massive spectroscopic stellar survey that observed almost half a million stars in the Galaxy. In a first study we use three different models of the Galaxy to generate synthetic stellar surveys that can be directly compared to the RAVE data. To do this we evaluate the RAVE selection function to great detail. Among the Galaxy models is the widely used Besancon model that performs well when individual parameter distribution are considered, but fails when we study chemodynamic correlations. The other two models are based on distributions of mass particles instead of analytical distribution functions. This is the first time that such models are converted to the space of observables and are compared to a stellar survey. We show that these models can be competitive and in some aspects superior to analytic models, because of their self-consistent dynamic history. In the case of a full cosmological simulation of disk galaxy formation we can recover features in the synthetic survey that relate to the known issues of the model and hence proof that our technique is sensitive to the global structure of the model. We argue that the next generation of cosmological galaxy formation simulations will deliver valuable models for our Galaxy. Testing these models with our approach will provide a direct connection between stellar Galactic astronomy and physical cosmology. In the second part of the thesis we use a sample of high-velocity halo stars from the RAVE data to estimate the Galactic escape speed and the virial mass of the Milky Way. In the course of this study cosmological simulations of galaxy formation also play a crucial role. Here we use them to calibrate and extensively test our analysis technique. We find the local Galactic escape speed to be 533 (+54/-41) km/s (90% confidence). With this result in combination with a simple mass model of the Galaxy we then construct an estimate of the virial mass of the Galaxy. For the mass profile of the dark matter halo we use two extreme models, a pure Navarro, Frenk & White (NFW) profile and an adiabatically contracted NFW profile. When we use statistics on the concentration parameter of these profile taken from large dissipationless cosmological simulations we obtain an estimate of the virial mass that is almost independent of the choice of the halo profile. For the mass M_340 enclosed within R_340 = 180 kpc we find 1.3 (+0.4/-0.3) x 10^12 M_sun. This value is in very good agreement with a number of other mass estimates in the literature that are based on independent data sets and analysis techniques. In the last part of this thesis we investigate a new possible channel to generate a population of Hypervelocity stars (HVSs) that is observed in the stellar halo. Commonly, it is assumed that the velocities of these stars originate from an interaction with the super-massive black hole in the Galactic center. It was suggested recently that stars stripped-off a disrupted satellite galaxy could reach similar velocities and leave the Galaxy. Here we study in detail the kinematics of tidal debris stars to investigate the probability that the observed sample of HVSs could partly originate from such a galaxy collision. We use a suite of $N$-body simulations following the encounter of a satellite galaxy with its Milky Way-type host galaxy. We quantify the typical pattern in angular and phase space formed by the debris stars and develop a simple model that predicts the kinematics of stripped-off stars. We show that the distribution of orbital energies in the tidal debris has a typical form that can be described quite accurately by a simple function. The main parameters determining the maximum energy kick a tidal debris star can get is the initial mass of the satellite and only to a lower extent its orbit. Main contributors to an unbound stellar population created in this way are massive satellites (M_sat > 10^9 M_sun). The probability that the observed HVS population is significantly contaminated by tidal debris stars appears small in the light of our results.
In the presence of a solid-liquid or liquid-air interface, bacteria can choose between a planktonic and a sessile lifestyle. Depending on environmental conditions, cells swimming in close proximity to the interface can irreversibly attach to the surface and grow into three-dimensional aggregates where the majority of cells is sessile and embedded in an extracellular polymer matrix (biofilm). We used microfluidic tools and time lapse microscopy to perform experiments with the polarly flagellated soil bacterium Pseudomonas putida (P. putida), a bacterial species that is able to form biofilms. We analyzed individual trajectories of swimming cells, both in the bulk fluid and in close proximity to a glass-liquid interface. Additionally, surface related growth during the early phase of biofilm formation was investigated. In the bulk fluid, P.putida shows a typical bacterial swimming pattern of alternating periods of persistent displacement along a line (runs) and fast reorientation events (turns) and cells swim with an average speed around 24 micrometer per second. We found that the distribution of turning angles is bimodal with a dominating peak around 180 degrees. In approximately six out of ten turning events, the cell reverses its swimming direction. In addition, our analysis revealed that upon a reversal, the cell systematically changes its swimming speed by a factor of two on average. Based on the experimentally observed values of mean runtime and rotational diffusion, we presented a model to describe the spreading of a population of cells by a run-reverse random walker with alternating speeds. We successfully recover the mean square displacement and, by an extended version of the model, also the negative dip in the directional autocorrelation function as observed in the experiments. The analytical solution of the model demonstrates that alternating speeds enhance a cells ability to explore its environment as compared to a bacterium moving at a constant intermediate speed. As compared to the bulk fluid, for cells swimming near a solid boundary we observed an increase in swimming speed at distances below d= 5 micrometer and an increase in average angular velocity at distances below d= 4 micrometer. While the average speed was maximal with an increase around 15% at a distance of d= 3 micrometer, the angular velocity was highest in closest proximity to the boundary at d=1 micrometer with an increase around 90% as compared to the bulk fluid. To investigate the swimming behavior in a confinement between two solid boundaries, we developed an experimental setup to acquire three-dimensional trajectories using a piezo driven objective mount coupled to a high speed camera. Results on speed and angular velocity were consistent with motility statistics in the presence of a single boundary. Additionally, an analysis of the probability density revealed that a majority of cells accumulated near the upper and lower boundaries of the microchannel. The increase in angular velocity is consistent with previous studies, where bacteria near a solid boundary were shown to swim on circular trajectories, an effect which can be attributed to a wall induced torque. The increase in speed at a distance of several times the size of the cell body, however, cannot be explained by existing theories which either consider the drag increase on cell body and flagellum near a boundary (resistive force theory) or model the swimming microorganism by a multipole expansion to account for the flow field interaction between cell and boundary. An accumulation of swimming bacteria near solid boundaries has been observed in similar experiments. Our results confirm that collisions with the surface play an important role and hydrodynamic interactions alone cannot explain the steady-state accumulation of cells near the channel walls. Furthermore, we monitored the number growth of cells in the microchannel under medium rich conditions. We observed that, after a lag time, initially isolated cells at the surface started to grow by division into colonies of increasing size, while coexisting with a comparable smaller number of swimming cells. After 5:50 hours, we observed a sudden jump in the number of swimming cells, which was accompanied by a breakup of bigger clusters on the surface. After approximately 30 minutes where planktonic cells dominated in the microchannel, individual swimming cells reattached to the surface. We interpret this process as an emigration and recolonization event. A number of complementary experiments were performed to investigate the influence of collective effects or a depletion of the growth medium on the transition. Similar to earlier observations on another bacterium from the same family we found that the release of cells to the swimming phase is most likely the result of an individual adaption process, where syntheses of proteins for flagellar motility are upregulated after a number of division cycles at the surface.
Organic semiconductors combine the benefits of organic materials, i.e., low-cost production, mechanical flexibility, lightweight, and robustness, with the fundamental semiconductor properties light absorption, emission, and electrical conductivity. This class of material has several advantages over conventional inorganic semiconductors that have led, for instance, to the commercialization of organic light-emitting diodes which can nowadays be found in the displays of TVs and smartphones. Moreover, organic semiconductors will possibly lead to new electronic applications which rely on the unique mechanical and electrical properties of these materials. In order to push the development and the success of organic semiconductors forward, it is essential to understand the fundamental processes in these materials. This thesis concentrates on understanding how the charge transport in thiophene-based semiconductor layers depends on the layer morphology and how the charge transport properties can be intentionally modified by doping these layers with a strong electron acceptor. By means of optical spectroscopy, the layer morphologies of poly(3-hexylthiophene), P3HT, P3HT-fullerene bulk heterojunction blends, and oligomeric polyquaterthiophene, oligo-PQT-12, are studied as a function of temperature, molecular weight, and processing conditions. The analyses rely on the decomposition of the absorption contributions from the ordered and the disordered parts of the layers. The ordered-phase spectra are analyzed using Spano’s model. It is figured out that the fraction of aggregated chains and the interconnectivity of these domains is fundamental to a high charge carrier mobility. In P3HT layers, such structures can be grown with high-molecular weight, long P3HT chains. Low and medium molecular weight P3HT layers do also contain a significant amount of chain aggregates with high intragrain mobility; however, intergranular connectivity and, therefore, efficient macroscopic charge transport are absent. In P3HT-fullerene blend layers, a highly crystalline morphology that favors the hole transport and the solar cell efficiency can be induced by annealing procedures and the choice of a high-boiling point processing solvent. Based on scanning near-field and polarization optical microscopy, the morphology of oligo-PQT-12 layers is found to be highly crystalline which explains the rather high field-effect mobility in this material as compared to low molecular weight polythiophene fractions. On the other hand, crystalline dislocations and grain boundaries are identified which clearly limit the charge carrier mobility in oligo-PQT-12 layers. The charge transport properties of organic semiconductors can be widely tuned by molecular doping. Indeed, molecular doping is a key to highly efficient organic light-emitting diodes and solar cells. Despite this vital role, it is still not understood how mobile charge carriers are induced into the bulk semiconductor upon the doping process. This thesis contains a detailed study of the doping mechanism and the electrical properties of P3HT layers which have been p-doped by the strong molecular acceptor tetrafluorotetracyanoquinodimethane, F4TCNQ. The density of doping-induced mobile holes, their mobility, and the electrical conductivity are characterized in a broad range of acceptor concentrations. A long-standing debate on the nature of the charge transfer between P3HT and F4TCNQ is resolved by showing that almost every F4TCNQ acceptor undergoes a full-electron charge transfer with a P3HT site. However, only 5% of these charge transfer pairs can dissociate and induce a mobile hole into P3HT which contributes electrical conduction. Moreover, it is shown that the left-behind F4TCNQ ions broaden the density-of-states distribution for the doping-induced mobile holes, which is due to the longrange Coulomb attraction in the low-permittivity organic semiconductors.
The problem under consideration in the thesis is a two level atom in a photonic crystal and a pumping laser. The photonic crystal provides an environment for the atom, that modifies the decay of the exited state, especially if the atom frequency is close to the band gap. The population inversion is investigated als well as the emission spectrum. The dynamics is analysed in the context of open quantum systems. Due to the multiple reflections in the photonic crystal, the system has a finite memory that inhibits the Markovian approximation. In the Heisenberg picture the equations of motion for the system variables form a infinite hierarchy of integro-differential equations. To get a closed system, approximations like a weak coupling approximation are needed. The thesis starts with a simple photonic crystal that is amenable to analytic calculations: a one-dimensional photonic crystal, that consists of alternating layers. The Bloch modes inside and the vacuum modes outside a finite crystal are linked with a transformation matrix that is interpreted as a transfer matrix. Formulas for the band structure, the reflection from a semi-infinite crystal, and the local density of states in absorbing crystals are found; defect modes and negative refraction are discussed. The quantum optics section of the work starts with the discussion of three problems, that are related to the full resonance fluorescence problem: a pure dephasing model, the driven atom and resonance fluorescence in free space. In the lowest order of the system-environment coupling, the one-time expectation values for the full problem are calculated analytically and the stationary states are discussed for certain cases. For the calculation of the two time correlation functions and spectra, the additional problem of correlations between the two times appears. In the Markovian case, the quantum regression theorem is valid. In the general case, the fluctuation dissipation theorem can be used instead. The two-time correlation functions are calculated by the two different methods. Within the chosen approximations, both methods deliver the same result. Several plots show the dependence of the spectrum on the parameters. Some examples for squeezing spectra are shown with different approximations. A projection operator method is used to establish two kinds of Markovian expansion with and without time convolution. The lowest order is identical with the lowest order of system environment coupling, but higher orders give different results.
The life of microorganisms is characterized by two main tasks, rapid growth under conditions permitting growth and survival under stressful conditions. The environments, in which microorganisms dwell, vary in space and time. The microorganisms innovate diverse strategies to readily adapt to the regularly fluctuating environments. Phenotypic heterogeneity is one such strategy, where an isogenic population splits into subpopulations that respond differently under identical environments. Bacterial persistence is a prime example of such phenotypic heterogeneity, whereby a population survives under an antibiotic attack, by keeping a fraction of population in a drug tolerant state, the persister state. Specifically, persister cells grow more slowly than normal cells under growth conditions, but survive longer under stress conditions such as the antibiotic administrations. Bacterial persistence is identified experimentally by examining the population survival upon an antibiotic treatment and the population resuscitation in a growth medium. The underlying population dynamics is explained with a two state model for reversible phenotype switching in a cell within the population. We study this existing model with a new theoretical approach and present analytical expressions for the time scale observed in population growth and resuscitation, that can be easily used to extract underlying model parameters of bacterial persistence. In addition, we recapitulate previously known results on the evolution of such structured population under periodically fluctuating environment using our simple approximation method. Using our analysis, we determine model parameters for Staphylococcus aureus population under several antibiotics and interpret the outcome of cross-drug treatment. Next, we consider the expansion of a population exhibiting phenotype switching in a spatially structured environment consisting of two growth permitting patches separated by an antibiotic patch. The dynamic interplay of growth, death and migration of cells in different patches leads to distinct regimes in population propagation speed as a function of migration rate. We map out the region in parameter space of phenotype switching and migration rate to observe the condition under which persistence is beneficial. Furthermore, we present an extended model that allows mutation from the two phenotypic states to a resistant state. We find that the presence of persister cells may enhance the probability of resistant mutation in a population. Using this model, we explain the experimental results showing the emergence of antibiotic resistance in a Staphylococcus aureus population upon tobramycin treatment. In summary, we identify several roles of bacterial persistence, such as help in spatial expansion, development of multidrug tolerance and emergence of antibiotic resistance. Our study provides a theoretical perspective on the dynamics of bacterial persistence in different environmental conditions. These results can be utilized to design further experiments, and to develop novel strategies to eradicate persistent infections.
In dieser Arbeit werden Konzepte für die Diagnostik der großskaligen Zirkulation in der Troposphäre und Stratosphäre entwickelt. Der Fokus liegt dabei auf dem Energiehaushalt, auf der Wellenausbreitung und auf der Interaktion der atmosphärischen Wellen mit dem Grundstrom. Die Konzepte werden hergeleitet, wobei eine neue Form des lokalen Eliassen-Palm-Flusses unter Einbeziehung der Feuchte eingeführt wird. Angewendet wird die Diagnostik dann auf den Reanalysedatensatz ERA-Interim und einen durch beobachtete Meerestemperatur- und Eisdaten angetriebenen Lauf des ECHAM6 Atmosphärenmodells. Die diagnostischen Werkzeuge zur Analyse der großskaligen Zirkulation sind einerseits nützlich, um das Verständnis der Dynamik des Klimasystems weiter zu fördern. Andererseits kann das gewonnene Verständnis des Zusammenhangs von Energiequellen und -senken sowie deren Verknüpfung mit synoptischen und planetaren Wellensystemen und dem resultierenden Antrieb des Grundstroms auch verwendet werden, um Klimamodelle auf die korrekte Wiedergabe dieser Beobachtungen zu prüfen. Hier zeigt sich, dass die Abweichungen im untersuchten ECHAM6-Modelllauf bezüglich des Energiehaushalts klein sind, jedoch teils starke Abweichungen bezüglich der Ausbreitung von atmosphärischen Wellen existieren. Planetare Wellen zeigen allgemein zu große Intensitäten in den Eliassen-Palm-Flüssen, während innerhalb der Strahlströme der oberen Troposphäre der Antrieb des Grundstroms durch synoptische Wellen verfälscht ist, da deren vertikale Ausbreitung gegenüber den Beobachtungen verschoben ist. Untersucht wird auch der Einfluss von arktischen Meereisänderungen ausgehend vom Bedeckungsminimum im August/September bis in den Winter. Es werden starke positive Temperaturanomalien festgestellt, welche an der Oberfläche am größten sind. Diese führen vor allem im Herbst zur Intensivierung von synoptischen Systemen in den arktischen Breiten, da die Stabilität der troposphärischen Schichtung verringert ist. Im darauffolgenden Winter stellen sich barotrope bis in die Stratosphäre reichende Änderungen der großskaligen Zirkulation ein, welche auf Meereisänderungen zurückzuführen sind. Der meridionale Druckgradient sinkt und führt so zu einem Muster ähnlich einer negativen Phase der arktischen Oszillation in der Troposphäre und einem geschwächten Polarwirbel in der Stratosphäre. Diese Zusammenhänge werden ebenfalls in einem ECHAM6-Modelllauf untersucht, wobei vor allem der Erwärmungstrend in der Arktis zu gering ist. Die großskaligen Veränderungen im Winter können zum Teil auch im Modelllauf festgestellt werden, jedoch zeigen sich insbesondere in der Stratosphäre Abweichungen für die Periode mit der geringsten Eisausdehnung. Die vertikale Ausbreitung planetarer Wellen von der Troposphäre in die Stratosphäre ist in ECHAM6 mit sehr großen Abweichungen wiedergegeben. Somit stellt die Wellenausbreitung insgesamt den größten in dieser Arbeit festgestellten Mangel in ECHAM6 dar.