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"Shanahan, Murray"
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The role of cortical oscillations in a spiking neural network model of the basal ganglia
Although brain oscillations involving the basal ganglia (BG) have been the target of extensive research, the main focus lies disproportionally on oscillations generated within the BG circuit rather than other sources, such as cortical areas. We remedy this here by investigating the influence of various cortical frequency bands on the intrinsic effective connectivity of the BG, as well as the role of the latter in regulating cortical behaviour. To do this, we construct a detailed neural model of the complete BG circuit based on fine-tuned spiking neurons, with both electrical and chemical synapses as well as short-term plasticity between structures. As a measure of effective connectivity, we estimate information transfer between nuclei by means of transfer entropy. Our model successfully reproduces firing and oscillatory behaviour found in both the healthy and Parkinsonian BG. We found that, indeed, effective connectivity changes dramatically for different cortical frequency bands and phase offsets, which are able to modulate (or even block) information flow in the three major BG pathways. In particular, alpha (8-12Hz) and beta (13-30Hz) oscillations activate the direct BG pathway, and favour the modulation of the indirect and hyper-direct pathways via the subthalamic nucleus-globus pallidus loop. In contrast, gamma (30-90Hz) frequencies block the information flow from the cortex completely through activation of the indirect pathway. Finally, below alpha, all pathways decay gradually and the system gives rise to spontaneous activity generated in the globus pallidus. Our results indicate the existence of a multimodal gating mechanism at the level of the BG that can be entirely controlled by cortical oscillations, and provide evidence for the hypothesis of cortically-entrained but locally-generated subthalamic beta activity. These two findings suggest new insights into the pathophysiology of specific BG disorders.
Journal Article
Activity in perceptual classification networks as a basis for human subjective time perception
Despite being a fundamental dimension of experience, how the human brain generates the perception of time remains unknown. Here, we provide a novel explanation for how human time perception might be accomplished, based on non-temporal perceptual classification processes. To demonstrate this proposal, we build an artificial neural system centred on a feed-forward image classification network, functionally similar to human visual processing. In this system, input videos of natural scenes drive changes in network activation, and accumulation of salient changes in activation are used to estimate duration. Estimates produced by this system match human reports made about the same videos, replicating key qualitative biases, including differentiating between scenes of walking around a busy city or sitting in a cafe or office. Our approach provides a working model of duration perception from stimulus to estimation and presents a new direction for examining the foundations of this central aspect of human experience.
How the brain tracks the passage of time remains unclear. Here, the authors show that tracking activation changes in a neural network trained to recognize objects (similar to the human visual system) produces estimates of duration that are subject to human-like biases.
Journal Article
Effects of lesions on synchrony and metastability in cortical networks
At the macroscopic scale, the human brain can be described as a complex network of white matter tracts integrating grey matter assemblies — the human connectome. The structure of the connectome, which is often described using graph theoretic approaches, can be used to model macroscopic brain function at low computational cost. Here, we use the Kuramoto model of coupled oscillators with time-delays, calibrated with respect to empirical functional MRI data, to study the relation between the structure of the connectome and two aspects of functional brain dynamics — synchrony, a measure of general coherence, and metastability, a measure of dynamical flexibility. Specifically, we investigate the relationship between the local structure of the connectome, quantified using graph theory, and the synchrony and metastability of the model's dynamics. By removing individual nodes and all of their connections from the model, we study the effect of lesions on both global and local dynamics. Of the nine nodal graph-theoretical properties tested, two were able to predict effects of node lesion on the global dynamics. The removal of nodes with high eigenvector centrality leads to decreases in global synchrony and increases in global metastability, as does the removal of hub nodes joining topologically segregated network modules. At the level of local dynamics in the neighbourhood of the lesioned node, structural properties of the lesioned nodes hold more predictive power, as five nodal graph theoretical measures are related to changes in local dynamics following node lesions. We discuss these results in the context of empirical studies of stroke and functional brain dynamics.
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•An oscillator model is coupled according to a human cortical structural connectome.•Model dynamics are calibrated to match empirical fMRI data.•Nodal properties of the connectome are quantified using graph theory.•Changes in synchrony and metastability following nodal lesions are quantified.•Lesions reveal relationships between connectome structure and modelled dynamics.
Journal Article
Correction: The role of cortical oscillations in a spiking neural network model of the basal ganglia
[This corrects the article DOI: 10.1371/journal.pone.0189109.].
Journal Article
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.
Journal Article
Role play with large language models
As dialogue agents become increasingly human-like in their performance, we must develop effective ways to describe their behaviour in high-level terms without falling into the trap of anthropomorphism. Here we foreground the concept of role play. Casting dialogue-agent behaviour in terms of role play allows us to draw on familiar folk psychological terms, without ascribing human characteristics to language models that they in fact lack. Two important cases of dialogue-agent behaviour are addressed this way, namely, (apparent) deception and (apparent) self-awareness.
By casting large-language-model-based dialogue-agent behaviour in terms of role play, it is possible to describe dialogue-agent behaviour such as (apparent) deception and (apparent) self-awareness without misleadingly ascribing human characteristics to the models.
Journal Article
The Partial Information Decomposition of Generative Neural Network Models
In this work we study the distributed representations learnt by generative neural network models. In particular, we investigate the properties of redundant and synergistic information that groups of hidden neurons contain about the target variable. To this end, we use an emerging branch of information theory called partial information decomposition (PID) and track the informational properties of the neurons through training. We find two differentiated phases during the training process: a first short phase in which the neurons learn redundant information about the target, and a second phase in which neurons start specialising and each of them learns unique information about the target. We also find that in smaller networks individual neurons learn more specific information about certain features of the input, suggesting that learning pressure can encourage disentangled representations.
Journal Article
Large-scale network organization in the avian forebrain: a connectivity matrix and theoretical analysis
Many species of birds, including pigeons, possess demonstrable cognitive capacities, and some are capable of cognitive feats matching those of apes. Since mammalian cortex is laminar while the avian telencephalon is nucleated, it is natural to ask whether the brains of these two cognitively capable taxa, despite their apparent anatomical dissimilarities, might exhibit common principles of organization on some level. Complementing recent investigations of macro-scale brain connectivity in mammals, including humans and macaques, we here present the first large-scale \"wiring diagram\" for the forebrain of a bird. Using graph theory, we show that the pigeon telencephalon is organized along similar lines to that of a mammal. Both are modular, small-world networks with a connective core of hub nodes that includes prefrontal-like and hippocampal structures. These hub nodes are, topologically speaking, the most central regions of the pigeon's brain, as well as being the most richly connected, implying a crucial role in information flow. Overall, our analysis suggests that indeed, despite the absence of cortical layers and close to 300 million years of separate evolution, the connectivity of the avian brain conforms to the same organizational principles as the mammalian brain.
Journal Article
The brain's connective core and its role in animal cognition
This paper addresses the question of how the brain of an animal achieves cognitive integration—that is to say how it manages to bring its fullest resources to bear on an ongoing situation. To fully exploit its cognitive resources, whether inherited or acquired through experience, it must be possible for unanticipated coalitions of brain processes to form. This facilitates the novel recombination of the elements of an existing behavioural repertoire, and thereby enables innovation. But in a system comprising massively many anatomically distributed assemblies of neurons, it is far from clear how such open-ended coalition formation is possible. The present paper draws on contemporary findings in brain connectivity and neurodynamics, as well as the literature of artificial intelligence, to outline a possible answer in terms of the brain's most richly connected and topologically central structures, its so-called connective core.
Journal Article