Recent developments in the quantitative analysis of complex networks, based largely on graph theory, have been rapidly translated to studies of brain network organization. The brain's structural and functional systems have features of complex networks--such as small-world topology, highly connected hubs and modularity--both at the whole-brain scale of human neuroimaging and at a cellular scale in non-human animals. In this article, we review studies investigating complex brain networks in diverse experimental modalities (including structural and functional MRI, diffusion tensor imaging, magnetoencephalography and electroencephalography in humans) and provide an accessible introduction to the basic principles of graph theory. We also highlight some of the technical challenges and key questions to be addressed by future developments in this rapidly moving field.

Complex brain networks: graph theoretical analysis of structural and functional systems

Graph theory is a valuable framework to study the organization of functional and anatomical connections in the brain. Its use for comparing network topologies, however, is not without difficulties. Graph measures may be influenced by the number of nodes (N) and the average degree (k) of the network. The explicit form of that influence depends on the type of network topology, which is usually unknown for experimental data. Direct comparisons of graph measures between empirical networks with different N and/or k can therefore yield spurious results. We list benefits and pitfalls of various approaches that intend to overcome these difficulties. We discuss the initial graph definition of unweighted graphs via fixed thresholds, average degrees or edge densities, and the use of weighted graphs. For instance, choosing a threshold to fix N and k does eliminate size and density effects but may lead to modifications of the network by enforcing (ignoring) non-significant (significant) connections. Opposed to fixing N and k, graph measures are often normalized via random surrogates but, in fact, this may even increase the sensitivity to differences in N and k for the commonly used clustering coefficient and small-world index. To avoid such a bias we tried to estimate the N,k-dependence for empirical networks, which can serve to correct for size effects, if successful. We also add a number of methods used in social sciences that build on statistics of local network structures including exponential random graph models and motif counting. We show that none of the here-investigated methods allows for a reliable and fully unbiased comparison, but some perform better than others.

/pdf/comparing-brain-networks-of-different-size-and-connectivity-173xg2dsac.pdf

Comparing brain networks of different size and connectivity density using graph theory

Brain graphs provide a relatively simple and increasingly popular way of modeling the human brain connectome, using graph theory to abstractly define a nervous system as a set of nodes (denoting anatomical regions or recording electrodes) and interconnecting edges (denoting structural or functional connections). Topological and geometrical properties of these graphs can be measured and compared to random graphs and to graphs derived from other neuroscience data or other (nonneural) complex systems. Both structural and functional human brain graphs have consistently demonstrated key topological properties such as small-worldness, modularity, and heterogeneous degree distributions. Brain graphs are also physically embedded so as to nearly minimize wiring cost, a key geometric property. Here we offer a conceptual review and methodological guide to graphical analysis of human neuroimaging data, with an emphasis on some of the key assumptions, issues, and trade-offs facing the investigator.

/pdf/brain-graphs-graphical-models-of-the-human-brain-connectome-135h0nfght.pdf

Brain Graphs: Graphical Models of the Human Brain Connectome

Purpose of review
Recent developments in the statistical physics of complex networks have been translated to neuroimaging data in an effort to enhance our understanding of human brain structural and functional networks. This review focuses on studies using graph theoretical measures applied to structural MRI, diffusion MRI, functional MRI, electroencephalography and magnetoencephalography data.

/pdf/human-brain-networks-in-health-and-disease-3t4uibbsbg.pdf

Human Brain Networks in Health and Disease

Purpose of review In recent years, there has been an explosion of studies on network modeling of brain connectivity. This review will focus mainly on recent findings concerning graph theoretical analysis of human brain networks with a variety of imaging modalities, including structural MRI, diffusion MRI, functional MRI, and EEG/MEG. Recent findings Recent studies have utilized graph theoretical approaches to investigate the organizational principles of brain networks. These studies have consistently shown many important statistical properties underlying the topological organization of the human brain, including modularity, small-worldness, and the existence of highly connected network hubs. Importantly, these quantifiable network properties were found to change during normal development, aging, and various neurological and neuropsychiatric diseases such as Alzheimer's disease and schizophrenia. Moreover, several studies have also suggested that these network properties correlate with behavioral and genetic factors. Summary The exciting research regarding graph theoretical analysis of brain connectivity yields truly integrative and comprehensive descriptions of the structural and functional organization of the human brain, which provides important implications for health and disease. Future research will most likely involve integrative models of brain structural and functional connectivity with multimodal neuroimaging data, exploring whether graph-based brain network analysis could yield reliable biomarkers for disease diagnosis and treatment.

Graph theoretical modeling of brain connectivity.

Objective:

To determine the functional connectivity of different EEG bands at the “baseline” situation (rest) and during mathematical thinking in children and young adults to study the maturation effect on brain networks at rest and during a cognitive task.



Methods:

Twenty children (8–12 years) and twenty students (21–26 years) were studied. The synchronization likelihood was used to evaluate the interregional synchronization of different EEG frequency bands in children and adults, at rest and during math. Then, graphs were constructed and characterized in terms of local structure (clustering coefficient) and overall integration (path length) and the “optimal” organization of the connectivity i.e., the small world network (SWN).



Results:

The main findings were: (i) Enhanced synchronization for theta band during math more prominent in adults. (ii) Decrease of the optimal SWN organization of the alpha2 band during math. (iii) The beta and especially gamma bands showed lower synchronization and signs of lower SWN organization in both situations in adults.



Conclusion:

There are interesting findings related to the two age groups and the two situations. The theta band showed higher synchronization during math in adults as a result of higher capacity of the working memory in this age group. The alpha2 band showed some SWN disorganization during math, a process analog to the known desynchronization. In adults, a dramatic reduction of the connections in gray matter occurs. Although this maturation process is probably related to higher efficiency, reduced connectivity is expressed by lower synchronization and lower mean values of the graph parameters in adults. Hum Brain Mapp 2009. © 2007 Wiley-Liss, Inc.

https://onlinelibrary.wiley.com/doi/pdf/10.1002/hbm.20492

Kassia Kanatsouli

Papers

The Influence of Ageing on Complex Brain Networks : A Graph Theoretical Analysis