The waking brain: an update

Cell Mol Life Sci. 2011 Aug;68(15):2499-512. doi: 10.1007/s00018-011-0631-8. Epub 2011 Feb 13.


Wakefulness and consciousness depend on perturbation of the cortical soliloquy. Ascending activation of the cerebral cortex is characteristic for both waking and paradoxical (REM) sleep. These evolutionary conserved activating systems build a network in the brainstem, midbrain, and diencephalon that contains the neurotransmitters and neuromodulators glutamate, histamine, acetylcholine, the catecholamines, serotonin, and some neuropeptides orchestrating the different behavioral states. Inhibition of these waking systems by GABAergic neurons allows sleep. Over the past decades, a prominent role became evident for the histaminergic and the orexinergic neurons as a hypothalamic waking center.

Publication types

  • Review

MeSH terms

  • Animals
  • Arousal / physiology
  • Biogenic Monoamines / metabolism
  • Biogenic Monoamines / physiology
  • Brain / physiology*
  • Histamine / metabolism
  • Histamine / physiology
  • Humans
  • Hypothalamus, Posterior / physiology
  • Models, Biological
  • Wakefulness / physiology*


  • Biogenic Monoamines
  • Histamine