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      Changes in global and regional modularity associated with increasing working memory load

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          Abstract

          Using graph theory measures common to complex network analyses of neuroimaging data, the objective of this study was to explore the effects of increasing working memory processing load on functional brain network topology in a cohort of young adults. Measures of modularity in complex brain networks quantify how well a network is organized into densely interconnected communities. We investigated changes in both the large-scale modular organization of the functional brain network as a whole and regional changes in modular organization as demands on working memory increased from n = 1 to n = 2 on the standard n-back task. We further investigated the relationship between modular properties across working memory load conditions and behavioral performance. Our results showed that regional modular organization within the default mode and working memory circuits significantly changed from 1-back to 2-back task conditions. However, the regional modular organization was not associated with behavioral performance. Global measures of modular organization did not change with working memory load but were associated with individual variability in behavioral performance. These findings indicate that regional and global network properties are modulated by different aspects of working memory under increasing load conditions. These findings highlight the importance of assessing multiple features of functional brain network topology at both global and regional scales rather than focusing on a single network property.

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          Most cited references69

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          Spurious but systematic correlations in functional connectivity MRI networks arise from subject motion.

          Here, we demonstrate that subject motion produces substantial changes in the timecourses of resting state functional connectivity MRI (rs-fcMRI) data despite compensatory spatial registration and regression of motion estimates from the data. These changes cause systematic but spurious correlation structures throughout the brain. Specifically, many long-distance correlations are decreased by subject motion, whereas many short-distance correlations are increased. These changes in rs-fcMRI correlations do not arise from, nor are they adequately countered by, some common functional connectivity processing steps. Two indices of data quality are proposed, and a simple method to reduce motion-related effects in rs-fcMRI analyses is demonstrated that should be flexibly implementable across a variety of software platforms. We demonstrate how application of this technique impacts our own data, modifying previous conclusions about brain development. These results suggest the need for greater care in dealing with subject motion, and the need to critically revisit previous rs-fcMRI work that may not have adequately controlled for effects of transient subject movements. Copyright © 2011 Elsevier Inc. All rights reserved.
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            Modularity and community structure in networks

            M. Newman (2006)
            Many networks of interest in the sciences, including a variety of social and biological networks, are found to divide naturally into communities or modules. The problem of detecting and characterizing this community structure has attracted considerable recent attention. One of the most sensitive detection methods is optimization of the quality function known as "modularity" over the possible divisions of a network, but direct application of this method using, for instance, simulated annealing is computationally costly. Here we show that the modularity can be reformulated in terms of the eigenvectors of a new characteristic matrix for the network, which we call the modularity matrix, and that this reformulation leads to a spectral algorithm for community detection that returns results of better quality than competing methods in noticeably shorter running times. We demonstrate the algorithm with applications to several network data sets.
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              Finding and evaluating community structure in networks

              We propose and study a set of algorithms for discovering community structure in networks -- natural divisions of network nodes into densely connected subgroups. Our algorithms all share two definitive features: first, they involve iterative removal of edges from the network to split it into communities, the edges removed being identified using one of a number of possible "betweenness" measures, and second, these measures are, crucially, recalculated after each removal. We also propose a measure for the strength of the community structure found by our algorithms, which gives us an objective metric for choosing the number of communities into which a network should be divided. We demonstrate that our algorithms are highly effective at discovering community structure in both computer-generated and real-world network data, and show how they can be used to shed light on the sometimes dauntingly complex structure of networked systems.
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                Author and article information

                Contributors
                Journal
                Front Hum Neurosci
                Front Hum Neurosci
                Front. Hum. Neurosci.
                Frontiers in Human Neuroscience
                Frontiers Media S.A.
                1662-5161
                01 December 2014
                2014
                : 8
                : 954
                Affiliations
                [1] 1Laboratory for Complex Brain Networks, Wake Forest University School of Medicine Winston-Salem, NC, USA
                [2] 2Department of Psychology, Wake Forest University Winston-Salem, NC, USA
                [3] 3Department of Radiology, Wake Forest University School of Medicine Winston-Salem, NC, USA
                Author notes

                Edited by: John J. Foxe, Albert Einstein College of Medicine, USA

                Reviewed by: Nelson Cowan, University of Missouri, USA; Alexander Stevens, Oregon Health and Science University, USA; Cherie L. Marvel, Johns Hopkins University, USA

                *Correspondence: Matthew L. Stanley, Laboratory for Complex Brain Networks, Wake Forest University School of Medicine, 2000 W. 1st Street, Suite 701, Winston-Salem, NC 27104, USA e-mail: stanleymattl9@ 123456gmail.com

                This article was submitted to the journal Frontiers in Human Neuroscience.

                Article
                10.3389/fnhum.2014.00954
                4249452
                25520639
                52da343e-f65a-44fc-8642-4629b2a59abe
                Copyright © 2014 Stanley, Dagenbach, Lyday, Burdette and Laurienti.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution and reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 28 August 2014
                : 10 November 2014
                Page count
                Figures: 5, Tables: 2, Equations: 3, References: 99, Pages: 14, Words: 12232
                Categories
                Neuroscience
                Original Research Article

                Neurosciences
                working memory,working memory load,n-back,graph theory,fmri,complex network,modularity
                Neurosciences
                working memory, working memory load, n-back, graph theory, fmri, complex network, modularity

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