Measuring information-transfer delays

In complex networks such as gene networks, traffic systems or brain circuits it is important to understand how long it takes for the different parts of the network to effectively influence one another. In the brain, for example, axonal delays between brain areas can amount to several tens of milliseconds, adding an intrinsic component to any timing-based processing of information. Inferring neural interaction delays is thus needed to interpret the information transfer revealed by any analysis of directed interactions across brain structures. However, a robust estimation of interaction delays from neural activity faces several challenges if modeling assumptions on interaction mechanisms are wrong or cannot be made. Here, we propose a robust estimator for neuronal interaction delays rooted in an information-theoretic framework, which allows a model-free exploration of interactions. In particular, we extend transfer entropy to account for delayed source-target interactions, while crucially retaining the conditioning on the embedded target state at the immediately previous time step. We prove that this particular extension is indeed guaranteed to identify interaction delays between two coupled systems and is the only relevant option in keeping with Wiener’s principle of causality. We demonstrate the performance of our approach in detecting interaction delays on finite data by numerical simulations of stochastic and deterministic processes, as well as on local field potential recordings. We also show the ability of the extended transfer entropy to detect the presence of multiple delays, as well as feedback loops. While evaluated on neuroscience data, we expect the estimator to be useful in other fields dealing with network dynamics

TRENTOOL: A Matlab open source toolbox to analyse information flow in time series data with transfer entropy

Background: Transfer entropy (TE) is a measure for the detection of directed interactions. Transfer entropy is an information theoretic implementation of Wiener's principle of observational causality. It offers an approach to the detection of neuronal interactions that is free of an explicit model of the interactions. Hence, it offers the power to analyze linear and nonlinear interactions alike. This allows for example the comprehensive analysis of directed interactions in neural networks at various levels of description. Here we present the open-source MATLAB toolbox TRENTOOL that allows the user to handle the considerable complexity of this measure and to validate the obtained results using non-parametrical statistical testing. We demonstrate the use of the toolbox and the performance of the algorithm on simulated data with nonlinear (quadratic) coupling and on local field potentials (LFP) recorded from the retina and the optic tectum of the turtle (Pseudemys scripta elegans) where a neuronal one-way connection is likely present. Results: In simulated data TE detected information flow in the simulated direction reliably with false positives not exceeding the rates expected under the null hypothesis. In the LFP data we found directed interactions from the retina to the tectum, despite the complicated signal transformations between these stages. No false positive interactions in the reverse directions were detected. Conclusions: TRENTOOL is an implementation of transfer entropy and mutual information analysis that aims to support the user in the application of this information theoretic measure. TRENTOOL is implemented as a MATLAB toolbox and available under an open source license (GPL v3). For the use with neural data TRENTOOL seamlessly integrates with the popular FieldTrip toolbox

Mean field modelling of human EEG: application to epilepsy

Aggregated electrical activity from brain regions recorded via an electroencephalogram (EEG), reveal that the brain is never at rest, producing a spectrum of ongoing oscillations that change as a result of different behavioural states and neurological conditions. In particular, this thesis focusses on pathological oscillations associated with absence seizures that typically affect 2–16 year old children. Investigation of the cellular and network mechanisms for absence seizures studies have implicated an abnormality in the cortical and thalamic activity in the generation of absence seizures, which have provided much insight to the potential cause of this disease. A number of competing hypotheses have been suggested, however the precise cause has yet to be determined. This work attempts to provide an explanation of these abnormal rhythms by considering a physiologically based, macroscopic continuum mean-field model of the brain's electrical activity. The methodology taken in this thesis is to assume that many of the physiological details of the involved brain structures can be aggregated into continuum state variables and parameters. The methodology has the advantage to indirectly encapsulate into state variables and parameters, many known physiological mechanisms underlying the genesis of epilepsy, which permits a reduction of the complexity of the problem. That is, a macroscopic description of the involved brain structures involved in epilepsy is taken and then by scanning the parameters of the model, identification of state changes in the system are made possible. Thus, this work demonstrates how changes in brain state as determined in EEG can be understood via dynamical state changes in the model providing an explanation of absence seizures. Furthermore, key observations from both the model and EEG data motivates a number of model reductions. These reductions provide approximate solutions of seizure oscillations and a better understanding of periodic oscillations arising from the involved brain regions. Local analysis of oscillations are performed by employing dynamical systems theory which provide necessary and sufficient conditions for their appearance. Finally local and global stability is then proved for the reduced model, for a reduced region in the parameter space. The results obtained in this thesis can be extended and suggestions are provided for future progress in this area