Towards a biologically annotated brain connectome.

The brain is a network of interleaved neural circuits. In modern connectomics, brain connectivity is typically encoded as a network of nodes and edges, abstracting away the rich biological detail of local neuronal populations. Yet biological annotations for network nodes - such as gene expression, cytoarchitecture, neurotransmitter receptors or intrinsic dynamics - can be readily measured and overlaid on network models. Here we review how connectomes can be represented and analysed as annotated networks. Annotated connectomes allow us to reconceptualize architectural features of networks and to relate the connection patterns of brain regions to their underlying biology. Emerging work demonstrates that annotated connectomes help to make more veridical models of brain network formation, neural dynamics and disease propagation. Finally, annotations can be used to infer entirely new inter-regional relationships and to construct new types of network that complement existing connectome representations. In summary, biologically annotated connectomes offer a compelling way to study neural wiring in concert with local biological features.