Shapley ratings in brain networks

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.