Establishing Communication between Neuronal Populations through Competitive Entrainment

The role of gamma frequency oscillation in neuronal interaction, and the relationship between oscillation and information transfer between neurons, has been the focus of much recent research. While the biological mechanisms responsible for gamma oscillation and the properties of resulting networks are well studied, the dynamics of changing phase coherence between oscillating neuronal populations are not well understood. To this end we develop a computational model of competitive selection between multiple stimuli, where the selection and transfer of population-encoded information arises from competition between converging stimuli to entrain a target population of neurons. Oscillation is generated by Pyramidal-Interneuronal Network Gamma through the action of recurrent synaptic connections between a locally connected network of excitatory and inhibitory neurons. Competition between stimuli is driven by differences in coherence of oscillation, while transmission of a single selected stimulus is enabled between generating and receiving neurons via Communication-through-Coherence. We explore the effect of varying synaptic parameters on the competitive transmission of stimuli over different neuron models, and identify a continuous region within the parameter space of the recurrent synaptic loop where inhibition-induced oscillation results in entrainment of target neurons. Within this optimal region we find that competition between stimuli of equal coherence results in model output that alternates between representation of the stimuli, in a manner strongly resembling well-known biological phenomena resulting from competitive stimulus selection such as binocular rivalry

Oscillatory dynamics as a mechanism of integration in complex networks of neurons

The large-scale integrative mechanisms of the brain, the means by which the activity of functionally segregated neuronal regions are combined, are not well understood. There is growing agreement that a flexible mechanism of integration must be present in order to support the myriad changing cognitive demands under which we are placed. Neuronal communication through phase-coherent oscillation stands as the prominent theory of cognitive integration. The work presented in this thesis explores the role of oscillation and synchronisation in the transfer and integration of information in the brain. It is first shown that complex metastable dynamics suitable for modelling phase-coherent neuronal synchronisation emerge from modularity in networks of delay and pulse-coupled oscillators. Within a restricted parameter regime these networks display a constantly changing set of partially synchronised states where some modules remain highly synchronised while others desynchronise. An examination of network phase dynamics shows increasing coherence with increasing connectivity between modules. The metastable chimera states that emerge from the activity of modular oscillator networks are demonstrated to be synchronous with a constant phase relationship as would be required of a mechanism of large-scale neural integration. A specific example of functional phase-coherent synchronisation within a spiking neural system is then developed. Competitive stimulus selection between converging population encoded stimuli is demonstrated through entrainment of oscillation in receiving neurons. The behaviour of the model is shown to be analogous to well-known competitive processes of stimulus selection such as binocular rivalry, matching key experimentally observed properties for the distribution and correlation of periods of entrainment under differing stimuli strength. Finally two new measures of network centrality, knotty-centrality and set betweenness centrality, are developed and applied to empirically derived human structural brain connectivity data. It is shown that human brain organisation exhibits a topologically central core network within a modular structure consistent with the generation of synchronous oscillation with functional phase dynamics

Knotty-Centrality: Finding the Connective Core of a Complex Network

A network measure called knotty-centrality is defined that quantifies the extent to which a given subset of a graph’s nodes constitutes a densely intra-connected topologically central connective core. Using this measure, the knotty centre of a network is defined as a sub-graph with maximal knotty-centrality. A heuristic algorithm for finding subsets of a network with high knotty-centrality is presented, and this is applied to previously published brain structural connectivity data for the cat and the human, as well as to a number of other networks. The cognitive implications of possessing a connective core with high knotty-centrality are briefly discussed

A biophysical model of dynamic balancing of excitation and inhibition in fast oscillatory large-scale networks

Over long timescales, neuronal dynamics can be robust to quite large perturbations, such as changes in white matter connectivity and grey matter structure through processes including learning, aging, development and certain disease processes. One possible explanation is that robust dynamics are facilitated by homeostatic mechanisms that can dynamically rebalance brain networks. In this study, we simulate a cortical brain network using the Wilson-Cowan neural mass model with conduction delays and noise, and use inhibitory synaptic plasticity (ISP) to dynamically achieve a spatially local balance between excitation and inhibition. Using MEG data from 55 subjects we find that ISP enables us to simultaneously achieve high correlation with multiple measures of functional connectivity, including amplitude envelope correlation and phase locking. Further, we find that ISP successfully achieves local E/I balance, and can consistently predict the functional connectivity computed from real MEG data, for a much wider range of model parameters than is possible with a model without ISP

The knotty centre of cat cortex and its relationship to other topologically significant subsets of nodes.

<p>There is good agreement in this case between rich club membership, high betweenness centrality, and knotty-centrality.</p

Example networks.

<p>(A) A network with a rich club. The set of nodes in the centre have high degree and are densely intra-connected. (B) A modular network with a knotty centre, but without a rich club. The set of nodes in the centre have high betweenness centrality, but their degree is no higher than the more peripheral nodes.</p