Discoveries over the past two decades demonstrate that regions distributed throughout the association cortex, often called the default network, are suppressed during tasks that demand external attention and are active during remembering, envisioning the future and making social inferences. This Review describes progress in understanding the organization and function of networks embedded within these association regions. Detailed high-resolution analyses of single individuals suggest that the default network is not a single network, as historically described, but instead comprises multiple interwoven networks. The multiple networks share a common organizational motif (also evident in marmoset and macaque anatomical circuits) that might support a general class of processing function dependent on internally constructed rather than externally constrained representations, with each separate interwoven network specialized for a distinct processing domain. Direct neuronal recordings in humans and monkeys reveal evidence for competitive relationships between the internally and externally oriented networks. Findings from rodent studies suggest that the thalamus might be essential to controlling which networks are engaged through specialized thalamic reticular neurons, including antagonistic subpopulations. These association networks (and presumably thalamocortical circuits) are expanded in humans and might be particularly vulnerable to dysregulation implicated in mental illness.
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The authors thank J. Andrews-Hanna and M. Halassa for helpful comments and R. Braga for discussion relating to this Review, and the reviewers who had extensive, constructive suggestions. D. Reznik helped re-plot the data from the UK Biobank. The marmoset tracer data were provided by the Marmoset Architecture Project. H. Becker assisted in preparation of the paper. The authors’ research work was supported by the US National Institutes of Health grant P50MH106435 to R.L.B. and the National Science Foundation Graduate Research Fellowship DGE1745303 to L.M.D.
The authors declare no competing interests.
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Human Connectome Project: https://www.humanconnectome.org/
Human Group-Averaged Network Estimates: http://freesurfer.net/fswiki/CorticalParcellation_Yeo2011
The Marmoset Architecture Project: www.marmosetbrain.org
UK Biobank: www.fmrib.ox.ac.uk/ukbiobank
- Task suppression
As used here, the reduced default network activity observed during an active task relative to its activity level during a passive (control) task.
- Intrinsic functional connectivity
Correlations between spatially separate brain regions in their spontaneous functional MRI activity signal that can be used to generate hypotheses about network organization.
Negative correlations in the spontaneous functional MRI signal that are present between separate networks of regions positively correlated among themselves.
- Internal mentation
Cognitive operations arising from internally constructed representations minimally dependent on stimuli in the immediate environment.
- Tract tracing
An anatomical method by which brain regions are injected to map the neurons receiving (anterograde) or sending (retrograde) projections from or to the region.
- Bowtie organization
As used here, refers to a core–periphery anatomical organization that resembles a bowtie with wings.
- High-frequency broadband
(HFB). High-frequency (>50 Hz) non-oscillatory activity recorded in the local field potential that reflects population-spiking activity.
- Thalamic reticular nucleus
(TRN). A modulatory nucleus surrounding the thalamus composed of GABAergic inhibitory neurons projecting to the thalamus.
- Sleep spindles
Abrupt bursts of oscillatory activity (12–14 Hz) generated by circuit interactions between the thalamic reticular nucleus and the thalamus during stage 2 sleep.
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Buckner, R.L., DiNicola, L.M. The brain’s default network: updated anatomy, physiology and evolving insights. Nat Rev Neurosci 20, 593–608 (2019). https://doi.org/10.1038/s41583-019-0212-7
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