Original Research ARTICLE

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Hierarchical modularity in human brain functional networks

1
Brain Mapping Unit, Department of Psychiatry, University of Cambridge, Cambridge, UK
2
Behavioural and Clinical Neurosciences Institute, University of Cambridge, Cambridge, UK
3
Institute for Mathematical Sciences, Imperial College, London, UK
4
Melbourne Neuropsychiatry Centre, Department of Psychiatry, University of Melbourne, VIC, Australia
5
GSK Clinical Unit Cambridge, Addenbrooke’s Hospital, Cambridge, UK
The idea that complex systems have a hierarchical modular organization originated in the early 1960s and has recently attracted fresh support from quantitative studies of large scale, real-life networks. Here we investigate the hierarchical modular (or “modules-within-modules”) decomposition of human brain functional networks, measured using functional magnetic resonance imaging in 18 healthy volunteers under no-task or resting conditions. We used a customized template to extract networks with more than 1800 regional nodes, and we applied a fast algorithm to identify nested modular structure at several hierarchical levels. We used mutual information, 0 < I < 1, to estimate the similarity of community structure of networks in different subjects, and to identify the individual network that is most representative of the group. Results show that human brain functional networks have a hierarchical modular organization with a fair degree of similarity between subjects, I = 0.63. The largest five modules at the highest level of the hierarchy were medial occipital, lateral occipital, central, parieto-frontal and fronto-temporal systems; occipital modules demonstrated less sub-modular organization than modules comprising regions of multimodal association cortex. Connector nodes and hubs, with a key role in inter-modular connectivity, were also concentrated in association cortical areas. We conclude that methods are available for hierarchical modular decomposition of large numbers of high resolution brain functional networks using computationally expedient algorithms. This could enable future investigations of Simon’s original hypothesis that hierarchy or near-decomposability of physical symbol systems is a critical design feature for their fast adaptivity to changing environmental conditions.
Keywords:
graph theory, brain, network, modularity, hierarchy, near-decomposability, information
Citation:
Meunier D, Lambiotte R, Fornito A, Ersche KD and Bullmore ET (2009). Hierarchical modularity in human brain functional networks. Front. Neuroinform. 3:37. doi: 10.3389/neuro.11.037.2009
Received:
20 March 2009;
 Paper pending published:
25 June 2009;
Accepted:
02 October 2009;
 Published online:
30 October 2009.

Edited by:

Marcus Kaiser, Newcastle University, UK

Reviewed by:

Roger Guimera, Northwestern University, USA
Pedro Valdes-Sosa, Cuban Neuroscience Center, Cuba
Copyright:
© 2009 Meunier, Lambiotte, Fornito, Ersche and Bullmore. This is an open-access article subject to an exclusive license agreement between the authors and the Frontiers Research Foundation, which permits unrestricted use, distribution, and reproduction in any medium, provided the original authors and source are credited.
*Correspondence:
Edward T. Bullmore, Brain Mapping Unit, Herchel Smith Building, Robinson Way, Cambridge CB2 0SZ, UK. e-mail: etb23@cam.ac.uk
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