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Die Arbeit befasst sich mit zwei funktionalen Grenzwertsätzen für skalierte Linienzählprozesse von anzestralen Selektionsgraphen. Dazu werden zwei Modelle aus der mathematischen Populationsgenetik betrachtet. Wir führen zuerst das Moran-Modell mit gerichteter Selektion mit konstanter Populationsgröße N in kontinuierlicher Zeit und den Linienzählprozess des anzestralen Selektionsgraphen (MASP) gemäß Krone und Neuhauser (Theor. Popul. Biol. 1997) ein. Die Hauptaussage dieser Abschlussarbeit besagt, dass der passend standardisierte MASP im Fall der moderaten Selektion für N gegen unendlich in Verteilung gegen einen Ornstein-Uhlenbeck-Prozess konvergiert. Das zweite betrachtete Modell ist das Cannings-Modell mit gerichteter Selektion in diskreter Zeit, das gemäß Boenkost, González Casanova, Pokalyuk und Wakolbinger (Electron. J. Probab. 2021) eingeführt wird. Für ein Teilregime der moderat schwachen Selektion wird bewiesen, dass die reskalierten Fluktuationen des Linienzählprozesses des anzestralen Selektionsgraphen im Cannings-Modell ebenfalls in Verteilung gegen einen Ornstein-Uhlenbeck-Prozess konvergieren.
Analysing survival or fixation probabilities for a beneficial allele is a prominent task in the field of theoretical population genetics. Haldane's asymptotics is an approximation for the fixation probability in the case of a single beneficial mutant with small selective advantage in a large population.
In this thesis we analyse the interplay between genetic drift and directional selection and prove Haldane's asymptotics in different settings: For the fixation probability in Cannings models with moderate selection and for the survival probability of a slightly supercritical branching processes in a random environment.
In Chapter 3 we introduce a class of Cannings models with selection that allow for a forward and backward construction. In particular, a Cannings ancestral selection process can be defined for this class of models, which counts the number of potential parents and is in sampling duality to the forward frequency process. By means of this duality the probability of fixation can be expressed through the expectation of the Cannings ancestral selection process in stationarity. A control of this expectation yields that the fixation probability fulfils Haldane's asymptotics in a regime of moderately weak selection (Thm. 8).
In Chapter 4 we study the fixation probability of Cannings models in a regime of moderately strong selection. Here couplings of the frequency process of beneficial individuals with slightly supercritical Galton-Watson processes imply that the fixation probability is given by Haldane's asymptotics (Thm. 9).
Lastly, in Chapter 5 we consider slightly supercritical branching processes in an independent and identically distributed random environment and study the probability of survival as the number of expected offspring tends from above to one. We show that only if variance and expectation of the random offspring mean are of the same order the random environment has a non-trivial influence on the probability of survival, which results in a modification of Haldane's asymptotics. Out of the critical parameter regime the population goes extinct or survives with a probability that fulfils Haldane's asymptotics (Thm. 10).
The proof establishes an expression for the survival probability in terms of the shape function of the random offspring generating functions. This expression exhibits similarities to perpetuities known from a financial context. Consequently, we prove a limiting theorem for perpetuities with vanishing interest rates (Thm. 11).
Precise timing of spikes between different neurons has been found to convey reliable information beyond the spike count. In contrast, the role of small phase delays with high temporal variability, as reported for example in oscillatory activity in the visual cortex, remains largely unclear. This issue becomes particularly important considering the high speed of neuronal information processing, which is assumed to be based on only a few milliseconds, or oscillation cycles within each processing step.
We investigate the role of small and imprecise phase delays with a stochastic spiking model that is strongly motivated by experimental observations. Within individual oscillation cycles the model contains only two signal parameters describing directly the rate and the phase. We specifically investigate two quantities, the probability of correct stimulus detection and the probability of correct change point detection, as a function of these signal parameters and within short periods of time such as individual oscillation cycles.
Optimal combinations of the signal parameters are derived that maximize these probabilities and enable comparison of pure rate, pure phase and combined codes. In particular, the gain in detection probability when adding imprecise phases to pure rate coding increases with the number of stimuli. More interestingly, imprecise phase delays can considerably improve the process of detecting changes in the stimulus, while also decreasing the probability of false alarms and thus, increasing robustness and speed of change point detection.
The results are applied to parameters extracted from empirical spike train recordings of neurons in the visual cortex in response to a number of visual stimuli. The results suggest that near-optimal combinations of rate and phase parameters can be implemented in the brain, and that phase parameters could particularly increase the quality of change point detection in cases of highly similar stimuli.
The thesis deals with the analysis and modeling of point processes emerging from different experiments in neuroscience. In particular, the description and detection of different types of variability changes in point processes is of interest.
A non-stationary rate or variance of life times is a well-known problem in the description of point processes like neuronal spike trains and can affect the results of further analyses requiring stationarity. Moreover, non-stationary parameters might also contain important information themselves. The goal of the first part of the thesis is the (further) development of a technique to detect both rate and variance changes that may occur in multiple time scales separately or simultaneously. A two-step procedure building on the multiple filter test (Messer et al., 2014) is used that first tests the null hypothesis of rate homogeneity allowing for an inhomogeneous variance and that estimates change points in the rate if the null hypothesis is rejected. In the second step, the null hypothesis of variance homogeneity is tested and variance change points are estimated. Rate change points are used as input. The main idea is the comparison of estimated variances in adjacent windows of different sizes sliding over the process. To determine the rejection threshold functionals of the Brownian motion are identified as limit processes under the null of variance homogeneity. The non-parametric procedure is not restricted to the case of at most one change point. It is shown in simulation studies that the corresponding test keeps the asymptotic significance level for a wide range of parameters and that the test power is remarkable. The practical applicability of the procedure is underlined by the analysis of neuronal spike trains.
Point processes resulting from experiments on bistable perception are analyzed in the second part of the thesis. Visual illusions allowing for than more possible perception lead to unpredictable changes of perception. In the thesis data from (Schmack et al., 2015) are used. A rotating sphere with switching perceived rotation direction was presented to the participants of the study. The stimulus was presented continuously and intermittently, i.e., with short periods of „blank display“ between the presentation periods. There are remarkable differences in the response patterns between the two types of presentation. During continuous presentation the distribution of dominance times, i.e., the intervals of constant perception, is a right-skewed and unimodal distribution with a mean of about five seconds. In contrast, during intermittent presentation one observes very long, stable dominance times of more than one minute interchanging with very short, unstable dominance times of less than five seconds, i.e., an increase of variability.
The main goal of the second part is to develop a model for the response patterns to bistable perception that builds a bridge between empirical data analysis and mechanistic modeling. Thus, the model should be able to describe both the response patterns to continuous presentation and to intermittent presentation. Moreover, the model should be fittable to typically short experimental data, and the model should allow for neuronal correlates. Current approaches often use detailed assumptions and large parameter sets, which complicate parameter estimation.
First, a Hidden Markov Model is applied. Second, to allow for neuronal correlates, a Hierarchical Brownian Model (HBM) is introduced, where perception is modeled by the competition of two neuronal populations. The activity difference between these two populations is described by a Brownian motion with drift fluctuating between two borders, where each first hitting time causes a perceptual change. To model the response patterns to intermittent presentation a second layer with competing neuronal populations (coding a stable and an unstable state) is assumed. Again, the data are described very well, and the hypothesis that the relative time in the stable state is identical in a group of patients with schizophrenia and a control group is rejected. To sum up, the HBM intends to link empirical data analysis and mechanistic modeling and provides interesting new hypotheses on potential neuronal mechanisms of cognitive phenomena.
We study exchangeable coalescent trees and the evolving genealogical trees in models for neutral haploid populations.
We show that every exchangeable infinite coalescent tree can be obtained as the genealogical tree of iid samples from a random marked metric measure space when the marks are added to the metric distances. We apply this representation to generalize the tree-valued Fleming-Viot process to include the case with dust in which the genealogical trees have isolated leaves.
Using the Donnelly-Kurtz lookdown approach, we describe all individuals ever alive in the population model by a random complete and separable metric space, the lookdown space, which we endow with a family of sampling measures. This yields a pathwise construction of tree-valued Fleming-Viot processes. In the case of coming down from infinity, we also read off a process whose state space is endowed with the Gromov-Hausdorff-Prohorov topology. This process has additional jumps at the extinction times of parts of the population.
In the case with only binary reproduction events, we construct the lookdown space also from the Aldous continuum random tree by removing the root and the highest leaf, and by deforming the metric in a way that corresponds to the time change that relates the Fleming-Viot process with a Dawson-Watanabe process. The sampling measures on the lookdown space are then image measures of the normalized local time measures.
We also show invariance principles for Markov chains that describe the evolving genealogy in Cannings models. For such Markov chains with values in the space of distance matrix distributions, we show convergence to tree-valued Fleming-Viot processes under the conditions of Möhle and Sagitov for the convergence of the genealogy at a fixed time to a coalescent with simultaneous multiple mergers. For the convergence of Markov chains with values in the space of marked metric measure spaces, an additional assumption is needed in the case with dust.
Die Populationsgenetik beschäftigt sich mit dem Einfluss von zufälliger Reproduktion, Rekombination, Migration, Mutation und Selektion auf die genetische Struktur einer Population.
In dieser Arbeit mit dem englischen Titel "Ancestral lines under mutation and selection" wird das Zusammenspiel von zufälliger Reproduktion, gerichteter Selektion und Zweiwegmutation untersucht.
Dazu betrachten wir eine haploide Population in der jedes Individuum zu jedem Zeitpunkt genau einen von zwei Typen aus S:={0,1} trägt. Dabei sei 1 der neutrale und 0 der selektiv bevorzugte Typ. Im Diffusionslimes sehr großer Populationen modellieren wir den Prozess der Frequenz der Typ-0-Individuen durch eine Wright-Fisher-Diffusion X:=(X_t) mit Mutation und gerichteter Selektion.
Zu jedem Zeitpunkt s gibt es genau ein Individuum, dessen Nachkommen ab einem bestimmten zukünftigen Zeitpunkt t>s die gesamte Population ausmachen werden. Wir nennen dieses Individuum den gemeinsamen Vorfahren zum Zeitpunkt s, da alle Individuen zu allen Zeitpunkten r>t von ihm abstammen. Sei R_{s} dessen Typ zum Zeitpunkt s. Wir nehmen an, dass der Prozess X zum Zeitpunkt 0 im Gleichgewicht ist und definieren die Wahrscheinlichkeit, dass der gemeinsame Vorfahre zum Zeitpunkt 0 Typ 0 hat, durch h(x):= P(R_{0}=0|X_{0}=x). Eine Darstellung von h(x) wurde bereits von Fearnhead (2002) und Taylor (2007) gefunden und dort mit vorwiegend analytischen Methoden bewiesen. In dieser Arbeit entwickeln wir in Kapitel 3 ein neues Teilchenbild, den pruned lookdown ancestral selection graph (pruned LD-ASG), der für sich selbst genommen interessant ist und eine neue probabilistische Interpretation der Darstellung von h(x) liefert.
Durch Erweiterung des Teilchenbildes auf Nachkommenverteilungen mit schweren Tails und mit Hilfe einer Siegmund Dualität gelingt es uns in Kapitel 4 das Resultat für h(x) von klassischen Wright-Fisher-Diffusionen auf Lambda-Wright-Fisher-Diffuison zu erweitern.
Eine Verbindung zwischen Ideen von Taylor (2007), der den gemeinsamen Prozess (X,R) untersucht hat, und einem von Fearnhead (2002) betrachteten Prozess (R,V), der die Entwicklung des Typs R des gemeinsamen Vorfahren in einer Umgebung von V sogenannten virtuellen Linien beschreibt, stellen wir in Kapitel 6 her. Wir bestimmen die gemeinsame Dynamik des Tripels (X,R,V). In Kapitel 7 betrachten wir ein diskretes Bild mit endlicher Populationsgröße N und schlagen dort eine Brücke zu Resultaten von Kluth, Hustedt und Baake (2013).
Des Weiteren entwickeln wir in Kapitel 5 dieser Arbeit einen Algorithmus zur Simulation der Typen einer Stichprobe von m Individuen, die aus einer Wright-Fisher-Population mit Mutation und Selektion im Gleichgewicht gezogen wird. Mittels dieses Algorithmus illustrieren wir die Typenverteilung für verschiedene Parameterwerte und Stichprobengrößen.
This work proposes to employ the (bursty) GLO model from Bingmer et. al (2011) to model the occurrence of tropical cyclones. We develop a Bayesian framework to estimate the parameters of the model and, particularly, employ a Markov chain Monte Carlo algorithm. This also allows us to develop a forecasting framework for future events.
Moreover, we assess the default probability of an insurance company that is exposed to claims that occur according to a GLO process and show that the model is able to substantially improve actuarial risk management if events occur in oscillatory bursts.
Optimierung von Phasen- und Ratenparametern in einem stochastischen Modell neuronaler Feueraktivität
(2014)
In unserem Gehirn wird Information von Neuronen durch die Emission von Spikes repräsentiert. Als wichtige Signalkomponenten werden hierbei die Rate (Anzahl Spikes), die Phase (zeitliche Verschiebung der Spikes) und synchrone Oszillationen (rhythmische Entladungen der Neuronen am selben Zyklus) diskutiert.
In dieser Arbeit wird untersucht, wie Rate und Phase für eine optimale Detektion miteinander kombiniert werden und abhängig vom gewählten Parameterbereich wird der Beitrag der Phase quantifiziert.
Dies wird anhand eines stochastischen Spiketrain-Modell untersucht, das hohe Ähnlichkeiten zu empirischen Spiketrains zeigt und die drei genannten Signalkomponenten beinhaltet. Das ELO-Modell („exponential lockig to a free oscillator“) ist in zwei Prozessstufen unterteilt: Im Hintergrund steht ein globaler Oszillationsprozess, der unabhängige und normal-verteilte Intervallabschnitte hervorbringt (Oszillation). An den Intervallgrenzen starten unabhängig, inhomogene Poisson-Prozesse (Synchronizität) mit exponentiell abnehmender Feuerrate, die durch eine stimulusspezifische Rate und Phase festgelegt ist.
Neben einer analytischen Bestimmung der optimalen Parameter im Falle reiner Raten- bzw. Phasencodierung, wird die gemeinsame Codierung anhand von Simulationsstudien analysiert.
A multiple filter test for the detection of rate changes in renewal processes with varying variance
(2014)
The thesis provides novel procedures in the statistical field of change point detection in time series.
Motivated by a variety of neuronal spike train patterns, a broad stochastic point process model is introduced. This model features points in time (change points), where the associated event rate changes. For purposes of change point detection, filtered derivative processes (MOSUM) are studied. Functional limit theorems for the filtered derivative processes are derived. These results are used to support novel procedures for change point detection; in particular, multiple filters (bandwidths) are applied simultaneously in oder to detect change points in different time scales.
The work presented in this thesis is devoted to two classes of mathematical population genetics models, namely the Kingman-coalescent and the Beta-coalescents. Chapters 2, 3 and 4 of the thesis include results concerned with the first model, whereas Chapter 5 presents contributions to the second class of models.