Human cortical excitability increases with time awake

Reto Huber, Hanna Mäki, Mario Rosanova, Silvia Casarotto, Paola Canali, Adenauer G. Casali, Giulio Tononi, Marcello Massimini

Research output: Contribution to journalArticlepeer-review


Prolonged wakefulness is associated not only with obvious changes in the way we feel and perform but also with well-known clinical effects, such as increased susceptibility to seizures, to hallucinations, and relief of depressive symptoms. These clinical effects suggest that prolonged wakefulness may be associated with significant changes in the state of cortical circuits. While recent animal experiments have reported a progressive increase of cortical excitability with time awake, no conclusive evidence could be gathered in humans. In this study, we combine transcranial magnetic stimulation (TMS) and electroencephalography (EEG) to monitor cortical excitability in healthy individuals as a function of time awake. We observed that the excitability of the human frontal cortex, measured as the immediate (0-20 ms) EEG reaction to TMS, progressively increases with time awake, from morning to evening and after one night of total sleep deprivation, and that it decreases after recovery sleep. By continuously monitoring vigilance, we also found that this modulation in cortical responsiveness is tonic and not attributable to transient fluctuations of the level of arousal. The present results provide noninvasive electrophysiological evidence that wakefulness is associated with a steady increase in the excitability of human cortical circuits that is rebalanced during sleep.

Original languageEnglish
Pages (from-to)332-338
Number of pages7
JournalCerebral Cortex
Issue number2
Publication statusPublished - Feb 2013


  • compensatory tracking task
  • EEG
  • human cortical excitability
  • sleep deprivation
  • transcranial magnetic stimulation

ASJC Scopus subject areas

  • Cognitive Neuroscience
  • Cellular and Molecular Neuroscience


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