A tutorial in connectome analysis: Topological and spatial features of brain networks

High-throughput methods for yielding the set of connections in a neural system, the connectome, are now being developed. This tutorial describes ways to analyze the topological and spatial organizations of the connectome at the macroscopic level of connectivity between brain regions as well as the microscopic level of connectivity between neurons. We will describe topological features at three different levels: the local scale of individual nodes, the regional scale of sets of nodes, and the global scale of the complete set of nodes in a network. Such features can be used to characterize components of a network and to compare different networks, e.g. the connectome of patients and control subjects for clinical studies. At the global scale, different types of networks can be distinguished and we will describe Erdös-Rényi random, scale-free, small-world, modular, and hierarchical archetypes of networks. Finally, the connectome also has a spatial organization and we describe methods for analyzing wiring lengths of neural systems. As an introduction for new researchers in the field of connectome analysis, we discuss the benefits and limitations of each analysis approach.