Neuromodulation of the mind-wandering brain state: the interaction between neuromodulatory tone, sharp wave-ripples and spontaneous thought.

Mind-wandering has become a captivating topic for cognitive neuroscientists. By now, it is reasonably well described in terms of its phenomenology and the large-scale neural networks that support it. However, we know very little about what neurobiological mechanisms trigger a mind-wandering episode and sustain the mind-wandering brain state. Here, we focus on the role of ascending neuromodulatory systems (i.e. acetylcholine, noradrenaline, serotonin and dopamine) in shaping mind-wandering. We advance the hypothesis that the hippocampal sharp wave-ripple (SWR) is a compelling candidate for a brain state that can trigger mind-wandering episodes. This hippocampal rhythm, which occurs spontaneously in quiescent behavioural states, is capable of propagating widespread activity in the default network and is functionally associated with recollective, associative, imagination and simulation processes. The occurrence of the SWR is heavily dependent on hippocampal neuromodulatory tone. We describe how the interplay of neuromodulators may promote the hippocampal SWR and trigger mind-wandering episodes. We then identify the global neuromodulatory signatures that shape the evolution of the mind-wandering brain state. Under our proposed framework, mind-wandering emerges due to the interplay between neuromodulatory systems that influence the transitions between brain states, which either facilitate, or impede, a wandering mind. This article is part of the theme issue 'Offline perception: voluntary and spontaneous perceptual experiences without matching external stimulation'.