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      Functional cartography of complex metabolic networks

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          Abstract

          High-throughput techniques are leading to an explosive growth in the size of biological databases and creating the opportunity to revolutionize our understanding of life and disease. Interpretation of these data remains, however, a major scientific challenge. Here, we propose a methodology that enables us to extract and display information contained in complex networks. Specifically, we demonstrate that one can (i) find functional modules in complex networks, and (ii) classify nodes into universal roles according to their pattern of intra- and inter-module connections. The method thus yields a ``cartographic representation'' of complex networks. Metabolic networks are among the most challenging biological networks and, arguably, the ones with more potential for immediate applicability. We use our method to analyze the metabolic networks of twelve organisms from three different super-kingdoms. We find that, typically, 80% of the nodes are only connected to other nodes within their respective modules, and that nodes with different roles are affected by different evolutionary constraints and pressures. Remarkably, we find that low-degree metabolites that connect different modules are more conserved than hubs whose links are mostly within a single module.

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          Most cited references15

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          Statistical mechanics of complex networks

          Complex networks describe a wide range of systems in nature and society, much quoted examples including the cell, a network of chemicals linked by chemical reactions, or the Internet, a network of routers and computers connected by physical links. While traditionally these systems were modeled as random graphs, it is increasingly recognized that the topology and evolution of real networks is governed by robust organizing principles. Here we review the recent advances in the field of complex networks, focusing on the statistical mechanics of network topology and dynamics. After reviewing the empirical data that motivated the recent interest in networks, we discuss the main models and analytical tools, covering random graphs, small-world and scale-free networks, as well as the interplay between topology and the network's robustness against failures and attacks.
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            Community structure in social and biological networks

            A number of recent studies have focused on the statistical properties of networked systems such as social networks and the World-Wide Web. Researchers have concentrated particularly on a few properties which seem to be common to many networks: the small-world property, power-law degree distributions, and network transitivity. In this paper, we highlight another property which is found in many networks, the property of community structure, in which network nodes are joined together in tightly-knit groups between which there are only looser connections. We propose a new method for detecting such communities, built around the idea of using centrality indices to find community boundaries. We test our method on computer generated and real-world graphs whose community structure is already known, and find that it detects this known structure with high sensitivity and reliability. We also apply the method to two networks whose community structure is not well-known - a collaboration network and a food web - and find that it detects significant and informative community divisions in both cases.
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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

                Journal
                2005-02-23
                Article
                10.1038/nature03288
                q-bio/0502035
                5ad795d1-4fa5-459b-a2d8-763fcf4d2774
                History
                Custom metadata
                Nature 433, 895-900 (2005)
                17 pages, 4 figures. Go to http://amaral.northwestern.edu for the PDF file of the reprint
                q-bio.MN cond-mat.dis-nn

                Molecular biology
                Molecular biology

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