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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.
In der Arbeit wird ein Testverfahren zum Prüfen der Varianzhomogenität der Lebenszeiten eines Erneuerungsprozesses entwickelt. Das Verfahren basiert auf der "Filtered-Derivative"-Methode. Zur Herleitung des Annahmebereichs werden zunächst Bootstrap-Permutationen genutzt, bevor zu einer asymptotischen Methode übergangen wird. Ein entsprechender funktionaler Grenzwertsatz wird skizziert. Aufbauend auf dem Test wird ein Multiple-Filter-Algorithmus zur genauen Detektion der Varianz-Change-Points besprochen. Schließlich folgt die Inklusion von vorher detektierten Ratenänderungen in das Verfahren. Der Test und der Algorithmus werden in Simulationsstudien evaluiert. Abschließend erfolgt eine Anwendung auf EEG-Daten.
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 stochastic model for the joint evaluation of burstiness and regularity in oscillatory spike trains
(2013)
The thesis provides a stochastic model to quantify and classify neuronal firing patterns of oscillatory spike trains. A spike train is a finite sequence of time points at which a neuron has an electric discharge (spike) which is recorded over a finite time interval. In this work, these spike times are analyzed regarding special firing patterns like the presence or absence of oscillatory activity and clusters (so called bursts). These bursts do not have a clear and unique definition in the literature. They are often fired in response to behaviorally relevant stimuli, e.g., an unexpected reward or a novel stimulus, but may also appear spontaneously. Oscillatory activity has been found to be related to complex information processing such as feature binding or figure ground segregation in the visual cortex. Thus, in the context of neurophysiology, it is important to quantify and classify these firing patterns and their change under certain experimental conditions like pharmacological treatment or genetical manipulation. In neuroscientific practice, the classification is often done by visual inspection criteria without giving reproducible results. Furthermore, descriptive methods are used for the quantification of spike trains without relating the extracted measures to properties of the underlying processes.
For that reason, a doubly stochastic point process model is proposed and termed 'Gaussian Locking to a free Oscillator' - GLO. The model has been developed on the basis of empirical observations in dopaminergic neurons and in cooperation with neurophysiologists. The GLO model uses as a first stage an unobservable oscillatory background rhythm which is represented by a stationary random walk whose increments are normally distributed. Two different model types are used to describe single spike firing or clusters of spikes. For both model types, the distribution of the random number of spikes per beat has different probability distributions (Bernoulli in the single spike case or Poisson in the cluster case). In the second stage, the random spike times are placed around their birth beat according to a normal distribution. These spike times represent the observed point process which has five easily interpretable parameters to describe the regularity and the burstiness of the firing patterns.
It turns out that the point process is stationary, simple and ergodic. It can be characterized as a cluster process and for the bursty firing mode as a Cox process. Furthermore, the distribution of the waiting times between spikes can be derived for some parameter combination. The conditional intensity function of the point process is derived which is also called autocorrelation function (ACF) in the neuroscience literature. This function arises by conditioning on a spike at time zero and measures the intensity of spikes x time units later. The autocorrelation histogram (ACH) is an estimate for the ACF. The parameters of the GLO are estimated by fitting the ACF to the ACH with a nonlinear least squares algorithm. This is a common procedure in neuroscientific practice and has the advantage that the GLO ACF can be computed for all parameter combinations and that its properties are closely related to the burstiness and regularity of the process. The precision of estimation is investigated for different scenarios using Monte-Carlo simulations and bootstrap methods.
The GLO provides the neuroscientist with objective and reproducible classification rules for the firing patterns on the basis of the model ACF. These rules are inspired by visual inspection criteria often used in neuroscientific practice and thus support and complement usual analysis of empirical spike trains. When applied to a sample data set, the model is able to detect significant changes in the regularity and burst behavior of the cells and provides confidence intervals for the parameter estimates.
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.
Zeitreihen von spontan auftretenden Topograpfien elektrischer Felder an der Kopfoberfläche, die durch eine Elektroenzephalografie (EEG) gemessen werden, zeigen Zeiträume („EEG-Microstates“), während denen die Topografie quasi-stabil ist. Diese EEG-Microstates werden üblicherweise dadurch analysiert, dass die zu spezifischen Zeitpunkten beobachteten Ausprägungen des EEGs in eine kleine Anzahl von prototypischen Topografien („Karten“) eingeteilt werden. Dadurch erhält man eine diskrete Kartensequenz.
Um die Struktur der Übergangswahrscheinlichkeiten in experimentellen Kartensequenzen zu beschreiben, werden diese Sequenzen durch eine reduzierte Markov-Kette modelliert mit nur einem Parameter pro Karte. Die Markov-Ketten können mithilfe von zwei bestimmten stochastischen Prozessen konstruiert werden. Durch den einen Prozess werden zufällige Intervalle definiert, die zufällig den verschiedenen Karten zugeordnet werden. Durch den anderen Prozess werden zufällige Abtastungszeitpunkte bestimmt, zu denen die Karte des jeweils aktuellen Intervalls abgelesen wird.
Neben der Motivation und Vorstellung des Markov-Ketten-Modells werden in dieser Arbeit Schätzer für die Modellparameter vorgeschlagen und diskutiert sowie ihre asymptotischen Varianzen hergeleitet. Zudem wird ein Anpassungstest durchgeführt und es werden Abwandlungen des Modells untersucht.
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.
Neuronal activity in the brain is often investigated in the presence of stimuli, termed externally driven activity. This stimulus-response-perspective has long been focussed on in order to find out how the nervous system responds to different stimuli. The neuronal response consists of baseline activity, so called spontaneous activity1, and activity which is caused by the stimulus. The baseline activity is often considered as constant over time which allows the identification of the stimulus-evoked part of the neuronal response by averaging over a set of trials.
However, during the last years it has been recognized that own dynamics of the nervous system plays an important role in information processing. As a consequence, spontaneous activity is no longer regarded only as background ’noise’ and its role in cortical processing is reconsidered. Therefore, the study of spontaneous firing pattern gains more importance as these patterns may shape neuronal responses to a larger extent as previously thought. For example, recent findings suggest that prestimulus activity can predict a person’s visual perception performance on a single trial basis (Hanslmayr et al., 2007). In this context, Ringach (2009) remarks that one can learn much about even the quiescent state of the brain which “underlies the importance of understanding cortical responses as the fusion of ongoing activity and sensory input”.
Taking into account that spontaneous activity reflects anything else but noise, new challenges arise when analysing neuronal data. In this thesis one of these problems related to the analysis of neuronal activity will be adressed, namely the nonstationarity of firing rates.
The present work consists of four chapters. First of all the introduction gives neurophysiological background information to get an idea of neuronal information processing. Afterwords the theory of point processes is provided which forms the basis for modeling neuronal spiking data. In the last section of the introduction a statement of the problem is given. Chapter 2 proposes an easily applicable statistical method for the detection of nonstationarity. It is applied to simulations and to real data in order to show its capabilities. Thereafter, four other approaches are presented which provide useful illustrations concerning the nonstationarity of the firing rate but share the problem that one cannot make objective statements on the basis of their results. They were developed in the course of establishing a suitable method. In chapter 4 the results are discussed and suggestions for further study are given.