Inferring a simple mechanism for alpha-blocking by fitting a neural population model to EEG spectra.

Alpha blocking, a phenomenon where the alpha rhythm is reduced by attention to a visual, auditory, tactile or cognitive stimulus, is one of the most prominent features of human electroencephalography (EEG) signals. Here we identify a simple physiological mechanism by which opening of the eyes causes attenuation of the alpha rhythm. We fit a neural population model to EEG spectra from 82 subjects, each showing a different degree of alpha blocking upon opening of their eyes. Though it has been notoriously difficult to estimate parameters by fitting such models, we show how, by regularizing the differences in parameter estimates between eyes-closed and eyes-open states, we can reduce the uncertainties in these differences without significantly compromising fit quality. From this emerges a parsimonious explanation for the spectral differences between states: Changes to just a single parameter, pei, corresponding to the strength of a tonic excitatory input to the inhibitory cortical population, are sufficient to explain the reduction in alpha rhythm upon opening of the eyes. We detect this by comparing the shift in each model parameter between eyes-closed and eyes-open states. Whereas changes in most parameters are weak or negligible and do not scale with the degree of alpha attenuation across subjects, the change in pei increases monotonically with the degree of alpha blocking observed. These results indicate that opening of the eyes reduces alpha activity by increasing external input to the inhibitory cortical population.

Parameter estimation and identifiability in a neural population model for electro-cortical activity.

Electroencephalography (EEG) provides a non-invasive measure of brain electrical activity. Neural population models, where large numbers of interacting neurons are considered collectively as a macroscopic system, have long been used to understand features in EEG signals. By tuning dozens of input parameters describing the excitatory and inhibitory neuron populations, these models can reproduce prominent features of the EEG such as the alpha-rhythm. However, the inverse problem, of directly estimating the parameters from fits to EEG data, remains unsolved. Solving this multi-parameter non-linear fitting problem will potentially provide a real-time method for characterizing average neuronal properties in human subjects. Here we perform unbiased fits of a 22-parameter neural population model to EEG data from 82 individuals, using both particle swarm optimization and Markov chain Monte Carlo sampling. We estimate how much is learned about individual parameters by computing Kullback-Leibler divergences between posterior and prior distributions for each parameter. Results indicate that only a single parameter, that determining the dynamics of inhibitory synaptic activity, is directly identifiable, while other parameters have large, though correlated, uncertainties. We show that the eigenvalues of the Fisher information matrix are roughly uniformly spaced over a log scale, indicating that the model is sloppy, like many of the regulatory network models in systems biology. These eigenvalues indicate that the system can be modeled with a low effective dimensionality, with inhibitory synaptic activity being prominent in driving system behavior.