Involvement of the default mode network under varying levels of cognitive effort.

Everyday cognitive functioning is characterized by constant alternations between different modes of information processing, driven by constant fluctuations in environmental demands. At the neural level, this is realized through corresponding dynamic shifts in functional activation and network connectivity. A distinction is often made between resting and task processing and between task-negative and task-positive functional networks. The Default Mode Network (DMN) is classically considered as a resting state (i.e. task-negative) network, upregulated in the absence of cognitive demands. In contrast, task-positive networks have been labelled the Extrinsic Mode Network (EMN). We investigated changes in brain activation and functional network connectivity in an experimental situation of repeated alterations between levels of cognitive effort, following a block-design. Using fMRI and a classic Stroop paradigm, participants switched back and forth between periods of no effort (resting), low effort (word reading, i.e. automatic processing based on learned internal representations and rules) and high effort (color naming, i.e. cognitively controlled perceptual processing of specific features of external stimuli). Results showed an expected EMN-activation for task versus resting contrasts, and DMN-activation for rest versus task contrasts. The DMN was in addition more strongly activated during periods of low effort contrasted with high effort, suggesting a gradual up- and down-regulation of the DMN network, depending on the level of demand and the type of processing required. The often reported "anti-correlation" between DMN and EMN was strongest during periods of low effort, indicating intermittent contributions of both networks. Taken together, these results challenge the traditional view of the DMN as solely a task-negative network. Instead, both the EMN and DMN may contribute to low-effort cognitive processing. In contrast, periods of resting and high effort are dominated by the DMN and EMN, respectively.