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Original Research Article
Shapley ratings in brain networks

1  Department of Cognitive Neuroscience, Radboud University Nijmegen Medical Centre, The Netherlands
2  Institute of Anatomy II, Heinrich Heine University, Germany
3  Vogt Brain Research Institute, Heinrich Heine University, Germany
4   Department of Computer Science, Heinrich Heine University, Germany
5  Department of Psychological and Brain Sciences, Indiana University, USA


Recent applications of network theory to brain networks as well as the expanding empirical databases of brain architecture spawn an interest in novel techniques for analyzing connectivity patterns in the brain. Treating individual brain structures as nodes in a directed graph model permits the application of graph theoretical concepts to the analysis of these structures within their large-scale connectivity networks. In this paper, we explore the application of concepts from graph and game theory toward this end. Specifically, we utilize the Shapley value principle, which assigns a rank to players in a coalition based upon their individual contributions to the collective profit of that coalition, to assess the contributions of individual brain structures to the graph derived from the global connectivity network. We report Shapley values for variations of a prefrontal network, as well as for a visual cortical network, which had both been extensively investigated previously. This analysis highlights particular nodes as strong or weak contributors to global connectivity. To understand the nature of their contribution, we compare the Shapley values obtained from these networks and appropriate controls to other previously described nodal measures of structural connectivity. We find a strong correlation between Shapley values and both betweenness centrality and connection density. Moreover, a stepwise multiple linear regression analysis indicates that approximately 79% of the variance in Shapley values obtained from random networks can be explained by betweenness centrality alone. Finally, we investigate the effects of local lesions on the Shapley ratings, showing that the present networks have an immense structural resistance to degradation. We discuss our results highlighting the use of such measures for characterizing the organization and functional role of brain networks.

Keywords: cerebral cortex, connectivity, game theory, graph analysis, neural network

Citation: Kötter R, Reid AT, Krumnack A, Wanke E and Sporns O (2007) Shapley ratings in brain networks. Front. Neuroinform. (2007) 1:2. doi:10.3389/neuro.11.002.2007

Received: 30 August 2007; paper pending published: 24 September 2007; accepted: 24 October 2007; published online: 30 November 2007.

Edited by: 
Jan G. Bjaalie, International Neuroinformatics Coordination Facility , Sweden; University of Oslo, Norway

Reviewed by: 
Gully A. Burns, USC Information Sciences Institute, USA
Petter Holme, Royal Institute Institute of Technology, Sweden

Copyright: © 2007 Kötter, Reid, Krumnack, Wanke and Sporns. 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: Rolf Kötter, Section Neurophysiology and Neuroinformatics, Department of Cognitive Neuroscience, Radboud University Nijmegen Medical Centre, POB 9101, 6500 HB Nijmegen, The Netherlands. e-mail: rk@cns.umcn.nl
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