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      Small-World Propensity and Weighted Brain Networks

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          Abstract

          Quantitative descriptions of network structure can provide fundamental insights into the function of interconnected complex systems. Small-world structure, diagnosed by high local clustering yet short average path length between any two nodes, promotes information flow in coupled systems, a key function that can differ across conditions or between groups. However, current techniques to quantify small-worldness are density dependent and neglect important features such as the strength of network connections, limiting their application in real-world systems. Here, we address both limitations with a novel metric called the Small-World Propensity (SWP). In its binary instantiation, the SWP provides an unbiased assessment of small-world structure in networks of varying densities. We extend this concept to the case of weighted brain networks by developing (i) a standardized procedure for generating weighted small-world networks, (ii) a weighted extension of the SWP, and (iii) a method for mapping observed brain network data onto the theoretical model. In applying these techniques to compare real-world brain networks, we uncover the surprising fact that the canonical biological small-world network, the C. elegans neuronal network, has strikingly low SWP. These metrics, models, and maps form a coherent toolbox for the assessment and comparison of architectural properties in brain networks.

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          Small-world brain networks.

          Many complex networks have a small-world topology characterized by dense local clustering or cliquishness of connections between neighboring nodes yet a short path length between any (distant) pair of nodes due to the existence of relatively few long-range connections. This is an attractive model for the organization of brain anatomical and functional networks because a small-world topology can support both segregated/specialized and distributed/integrated information processing. Moreover, small-world networks are economical, tending to minimize wiring costs while supporting high dynamical complexity. The authors introduce some of the key mathematical concepts in graph theory required for small-world analysis and review how these methods have been applied to quantification of cortical connectivity matrices derived from anatomical tract-tracing studies in the macaque monkey and the cat. The evolution of small-world networks is discussed in terms of a selection pressure to deliver cost-effective information-processing systems. The authors illustrate how these techniques and concepts are increasingly being applied to the analysis of human brain functional networks derived from electroencephalography/magnetoencephalography and fMRI experiments. Finally, the authors consider the relevance of small-world models for understanding the emergence of complex behaviors and the resilience of brain systems to pathological attack by disease or aberrant development. They conclude that small-world models provide a powerful and versatile approach to understanding the structure and function of human brain systems.
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            Uncovering the overlapping community structure of complex networks in nature and society

            Many complex systems in nature and society can be described in terms of networks capturing the intricate web of connections among the units they are made of. A key question is how to interpret the global organization of such networks as the coexistence of their structural subunits (communities) associated with more highly interconnected parts. Identifying these a priori unknown building blocks (such as functionally related proteins, industrial sectors and groups of people) is crucial to the understanding of the structural and functional properties of networks. The existing deterministic methods used for large networks find separated communities, whereas most of the actual networks are made of highly overlapping cohesive groups of nodes. Here we introduce an approach to analysing the main statistical features of the interwoven sets of overlapping communities that makes a step towards uncovering the modular structure of complex systems. After defining a set of new characteristic quantities for the statistics of communities, we apply an efficient technique for exploring overlapping communities on a large scale. We find that overlaps are significant, and the distributions we introduce reveal universal features of networks. Our studies of collaboration, word-association and protein interaction graphs show that the web of communities has non-trivial correlations and specific scaling properties.
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              Weak pairwise correlations imply strongly correlated network states in a neural population

              Biological networks have so many possible states that exhaustive sampling is impossible. Successful analysis thus depends on simplifying hypotheses, but experiments on many systems hint that complicated, higher order interactions among large groups of elements play an important role. In the vertebrate retina, we show that weak correlations between pairs of neurons coexist with strongly collective behavior in the responses of ten or more neurons. Surprisingly, we find that this collective behavior is described quantitatively by models that capture the observed pairwise correlations but assume no higher order interactions. These maximum entropy models are equivalent to Ising models, and predict that larger networks are completely dominated by correlation effects. This suggests that the neural code has associative or error-correcting properties, and we provide preliminary evidence for such behavior. As a first test for the generality of these ideas, we show that similar results are obtained from networks of cultured cortical neurons.
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                Author and article information

                Journal
                Sci Rep
                Sci Rep
                Scientific Reports
                Nature Publishing Group
                2045-2322
                25 February 2016
                2016
                : 6
                : 22057
                Affiliations
                [1 ]Department of Bioengineering, University of Pennsylvania , Philadelphia, PA 19104 USA
                [2 ]US Army Research Laboratory, Aberdeen Proving Ground , MD 21005, USA
                [3 ]Department of Mathematics and Computational and Data-Enabled Science and Engineering Program, University at Buffalo , Buffalo, NY 14260 USA
                [4 ]Department of Biomedical Engineering, Johns Hopkins University , Baltimore, MD 21218 USA
                [5 ]Department of Electrical and Systems Engineering, University of Pennsylvania , Philadelphia, PA 19104 USA
                Author notes
                Article
                srep22057
                10.1038/srep22057
                4766852
                26912196
                d862646f-6b7a-4c1a-8dfd-f2333a093204
                Copyright © 2016, Macmillan Publishers Limited

                This work is licensed under a Creative Commons Attribution 4.0 International License. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in the credit line; if the material is not included under the Creative Commons license, users will need to obtain permission from the license holder to reproduce the material. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/

                History
                : 23 June 2015
                : 03 February 2016
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