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Wakefulness

Neuroscience · NLM MeSH · updated 2026-10-29 · version 1

Citations verified against PubMedNot yet clinician-reviewed10 referencesApproximately 4 min read

A state in which there is an enhanced potential for sensitivity and an efficient responsiveness to external stimuli.

Table of Contents

1. Definition2. Research landscape3. Reading the evidence4. Open-access studies5. Key reviews6. References7. About this article

Definition and research guide

This page combines a MeSH definition, publication statistics and selected research abstracts. It is a starting point for reading the literature, not a medical textbook or a systematic evidence review.

Content available: 4 complete abstracts (767 words), 10 references. Full article text is not stored or reproduced here; follow the PMID, DOI or PMC links to read the original sources and their access terms.

Definition

A state in which there is an enhanced potential for sensitivity and an efficient responsiveness to external stimuli.

Definition: NLM Medical Subject Headings (MeSH) scope note (Public domain (U.S. National Library of Medicine); attribution requested). View the MeSH record.

Also known as: Wakefulnesses.

Where it fits in the MeSH classification

Broader topics
Arousal
See also
Sleep Initiation and Maintenance Disorders

Research landscape

PubMed indexes 22,134 records tagged with the MeSH heading “Wakefulness” (counts retrieved 2026-10-29). 4,175 of them (18.9%) were published from 2020 onward, and the busiest period so far is through 1999 with 6,905 records.

Reviews account for 1,983 records (9.0%) and meta-analyses for 126. Randomized controlled trials number 928 (4.2%) as a share of all records on the topic.

PubMed records by publication periodthrough 1999: 6,905 records; 2000-2009: 4,774 records; 2010-2019: 6,280 records; 2020 onward: 4,175 records6,905through 19994,7742000-20096,2802010-20194,1752020 onward
Records in PubMed indexed with this heading, by publication period. Real counts from NCBI E-utilities; the final period covers only part of the current year. Volume reflects how much has been published, not how strong the evidence is.
Show the data
PeriodRecords
through 19996,905
2000-20094,774
2010-20196,280
2020 onward4,175
Share of records by publication typeReviews: 9.0 percent; Meta-analyses: 0.6 percent; Randomized controlled trials: 4.2 percentReviews9.0% (1,983)Meta-analyses0.6% (126)Randomized controlled trials4.2% (928)
Share of all PubMed records on this heading that carry each publication type. Types overlap with other records, so the shares do not sum to 100%.
Publication-type chart data
Publication types among 22,134 PubMed records
TypeRecordsShare
Reviews1,9839.0%
Meta-analyses1260.6%
Randomized controlled trials9284.2%

Method: PubMed query "Wakefulness"[MeSH Terms] with publication-type and date filters, retrieved 2026-10-29.

Reading the evidence in neuroscience

Neuroscience draws on tools with different strengths: cellular and animal work can manipulate circuits directly, while human imaging and electrophysiology observe the living brain at lower resolution. Small imaging studies are prone to false positives, and findings in animals do not always carry over to people. Clinical neurology and psychiatry rely on trials and cohort studies whose outcomes are often symptom scales. Check the species, the sample size, how the outcome was measured and whether results were replicated.

Evidence from open-access studies

These open-access studies come from PubMed’s best-match results for this heading (papers whose main topic it is), ranked here by citation count, with one recent addition. Each abstract is reproduced in full under its recorded open licence so that you can read the authors’ own words. They illustrate the literature; they are not a systematic summary of it. Citation counts measure attention, not reliability or clinical benefit.

Study comparison — selected abstracts, not a systematic review
Study / authorsYearSource label / designEurope PMC citationsLicence / source
Associations between light exposure and sleep timing and sleepiness while awake in a sample of UK adults in everyday life
Didikoglu A, Mohammadian N, Johnson S, van Tongeren M, Wright P, Casson AJ, et al.
2023Research article
Design not recorded in structured metadata
62cc by
PMID 37812713 · DOI · PMC10589638
Wake-like skin patterning and neural activity during octopus sleep
Pophale A, Shimizu K, Mano T, Iglesias TL, Martin K, Hiroi M, et al.
2023Research article
Design not recorded in structured metadata
51cc by
PMID 37380770 · DOI · PMC10322707
Waves traveling over a map of visual space can ignite short-term predictions of sensory input
Benigno GB, Budzinski RC, Davis ZW, Reynolds JH, Muller L
2023Research article
Design not recorded in structured metadata
39cc by
PMID 37296131 · DOI · PMC10256723
Wake-promoting neuromodulators in Alzheimer's disease: Implications for sleep and brain clearance
Pedersen TJ, Clemens SG, Winer JR, Sharma A, Falgàs N, Lucey BP, et al.
2026Review article
Design not recorded in structured metadata
2cc by
PMID 41830117 · DOI · PMC13093618

Associations between light exposure and sleep timing and sleepiness while awake in a sample of UK adults in everyday life [1]

Research article Didikoglu A, Mohammadian N, Johnson S, van Tongeren M, Wright P, Casson AJ, et al. · · 2023 · cited 62 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Experimental and interventional studies show that light can regulate sleep timing and sleepiness while awake by setting the phase of circadian rhythms and supporting alertness.
Closing excerpt
Our study establishes a method for collecting longitudinal sleep and health/performance data in everyday life and provides evidence of associations between light exposure and important determinants of sleep health and performance.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (246 words)

Experimental and interventional studies show that light can regulate sleep timing and sleepiness while awake by setting the phase of circadian rhythms and supporting alertness. The extent to which differences in light exposure explain variations in sleep and sleepiness within and between individuals in everyday life remains less clear. Here, we establish a method to address this deficit, incorporating an open-source wearable wrist-worn light logger (SpectraWear) and smartphone-based online data collection. We use it to simultaneously record longitudinal light exposure (in melanopic equivalent daylight illuminance), sleep timing, and subjective alertness over seven days in a convenience sample of 59 UK adults without externally imposed circadian challenge (e.g., shift work or jetlag). Participants reliably had strong daily rhythms in light exposure but frequently were exposed to less light during the daytime and more light in pre-bedtime and sleep episodes than recommended [T. M. Brown et al., PLoS Biol. 20, e3001571 (2022)]. Prior light exposure over several hours was associated with lower subjective sleepiness with, in particular, brighter light in the late sleep episode and after wake linked to reduced early morning sleepiness (sleep inertia). Higher pre-bedtime light exposure was associated with longer sleep onset latency. Early sleep timing was correlated with more reproducible and robust daily patterns of light exposure and higher daytime/lower night-time light exposure. Our study establishes a method for collecting longitudinal sleep and health/performance data in everyday life and provides evidence of associations between light exposure and important determinants of sleep health and performance.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 37812713 · DOI · PMC10589638

Wake-like skin patterning and neural activity during octopus sleep [2]

Research article Pophale A, Shimizu K, Mano T, Iglesias TL, Martin K, Hiroi M, et al. · · 2023 · cited 51 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
While sleeping, many vertebrate groups alternate between at least two sleep stages: rapid eye movement and slow wave sleep1-4, in part characterized by wake-like and synchronous brain activity, respectively.
Closing excerpt
The range of similarities with vertebrates indicates that aspects of two-stage sleep in octopuses may represent convergent features of complex cognition.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (229 words)

While sleeping, many vertebrate groups alternate between at least two sleep stages: rapid eye movement and slow wave sleep1-4, in part characterized by wake-like and synchronous brain activity, respectively. Here we delineate neural and behavioural correlates of two stages of sleep in octopuses, marine invertebrates that evolutionarily diverged from vertebrates roughly 550 million years ago (ref. 5) and have independently evolved large brains and behavioural sophistication. 'Quiet' sleep in octopuses is rhythmically interrupted by approximately 60-s bouts of pronounced body movements and rapid changes in skin patterning and texture6. We show that these bouts are homeostatically regulated, rapidly reversible and come with increased arousal threshold, representing a distinct 'active' sleep stage. Computational analysis of active sleep skin patterning reveals diverse dynamics through a set of patterns conserved across octopuses and strongly resembling those seen while awake. High-density electrophysiological recordings from the central brain reveal that the local field potential (LFP) activity during active sleep resembles that of waking. LFP activity differs across brain regions, with the strongest activity during active sleep seen in the superior frontal and vertical lobes, anatomically connected regions associated with learning and memory function7-10. During quiet sleep, these regions are relatively silent but generate LFP oscillations resembling mammalian sleep spindles11,12 in frequency and duration. The range of similarities with vertebrates indicates that aspects of two-stage sleep in octopuses may represent convergent features of complex cognition.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 37380770 · DOI · PMC10322707

Waves traveling over a map of visual space can ignite short-term predictions of sensory input [3]

Research article Benigno GB, Budzinski RC, Davis ZW, Reynolds JH, Muller L · · 2023 · cited 39 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Recent analyses have found waves of neural activity traveling across entire visual cortical areas in awake animals.
Closing excerpt
These results suggest traveling waves may play an essential computational role in the visual system by embedding continuous spatiotemporal structures over spatial maps.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (151 words)

Recent analyses have found waves of neural activity traveling across entire visual cortical areas in awake animals. These traveling waves modulate the excitability of local networks and perceptual sensitivity. The general computational role of these spatiotemporal patterns in the visual system, however, remains unclear. Here, we hypothesize that traveling waves endow the visual system with the capacity to predict complex and naturalistic inputs. We present a network model whose connections can be rapidly and efficiently trained to predict individual natural movies. After training, a few input frames from a movie trigger complex wave patterns that drive accurate predictions many frames into the future solely from the network's connections. When the recurrent connections that drive waves are randomly shuffled, both traveling waves and the ability to predict are eliminated. These results suggest traveling waves may play an essential computational role in the visual system by embedding continuous spatiotemporal structures over spatial maps.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 37296131 · DOI · PMC10256723

Wake-promoting neuromodulators in Alzheimer's disease: Implications for sleep and brain clearance [4]

Review article Pedersen TJ, Clemens SG, Winer JR, Sharma A, Falgàs N, Lucey BP, et al. · · 2026 · cited 2 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Neuromodulatory subcortical systems (NSS) regulate arousal, cognition, and sleep-wake transitions through widespread influence on cortical and subcortical networks.
Closing excerpt
We propose that NSS dysfunction may drive a self-reinforcing cycle of disrupted sleep, impaired glymphatic clearance, and neurodegeneration, underscoring the need to better understand this relationship to inform pharmacological interventions slowing or preventing AD.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (141 words)

Neuromodulatory subcortical systems (NSS) regulate arousal, cognition, and sleep-wake transitions through widespread influence on cortical and subcortical networks. Increasing evidence links dysfunction of these systems to the pathogenesis of Alzheimer's disease (AD). Degeneration and dysregulation of NSS occurs during the preclinical phase of AD, preceding the onset of cognitive decline. Alterations of NSS are implicated in sleep disruption and impair sleep-dependent cerebrospinal fluid (CSF) clearance via the glymphatic system, a process involved in removing amyloid beta and other neurotoxic proteins. This review synthesizes current evidence linking wake-promoting neuromodulators-norepinephrine, histamine, and orexin-to AD pathology, with an emphasis on their convergent effects on sleep regulation and brain fluid dynamics. We propose that NSS dysfunction may drive a self-reinforcing cycle of disrupted sleep, impaired glymphatic clearance, and neurodegeneration, underscoring the need to better understand this relationship to inform pharmacological interventions slowing or preventing AD.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 41830117 · DOI · PMC13093618

Key reviews and meta-analyses

The most cited review-type papers among PubMed’s best-match results for this heading, listed with bibliographic details only. Follow the links for the full text and access terms.

  • Research article Epidemiology of the human circadian clock [5]
    Roenneberg T, Kuehnle T, Juda M, Kantermann T, Allebrandt K, Gordijn M, et al. · Sleep Med Rev · 2007 · cited 1,072 times (Europe PMC)
  • Research article Control of sleep and wakefulness [6]
    Brown RE, Basheer R, McKenna JT, Strecker RE, McCarley RW · Physiol Rev · 2012 · cited 1,059 times (Europe PMC)
  • Research article Sleep state switching [7]
    Saper CB, Fuller PM, Pedersen NP, Lu J, Scammell TE · Neuron · 2010 · cited 1,023 times (Europe PMC)
  • Research article International consensus on the assessment of bruxism: Report of a work in progress [8]
    Lobbezoo F, Ahlberg J, Raphael KG, Wetselaar P, Glaros AG, Kato T, et al. · J Oral Rehabil · 2018 · cited 937 times (Europe PMC)
  • Research article The influence of sleep quality, sleep duration and sleepiness on school performance in children and adolescents: A meta-analytic review [9]
    Dewald JF, Meijer AM, Oort FJ, Kerkhof GA, Bögels SM · Sleep Med Rev · 2010 · cited 881 times (Europe PMC)
  • Research article Neural Circuitry of Wakefulness and Sleep [10]
    Scammell TE, Arrigoni E, Lipton JO · Neuron · 2017 · cited 806 times (Europe PMC)

References

  1. Didikoglu A, Mohammadian N, Johnson S, van Tongeren M, Wright P, Casson AJ, et al.. Associations between light exposure and sleep timing and sleepiness while awake in a sample of UK adults in everyday life. . 2023. PMID 37812713 · DOI · PMC10589638
  2. Pophale A, Shimizu K, Mano T, Iglesias TL, Martin K, Hiroi M, et al.. Wake-like skin patterning and neural activity during octopus sleep. . 2023. PMID 37380770 · DOI · PMC10322707
  3. Benigno GB, Budzinski RC, Davis ZW, Reynolds JH, Muller L. Waves traveling over a map of visual space can ignite short-term predictions of sensory input. . 2023. PMID 37296131 · DOI · PMC10256723
  4. Pedersen TJ, Clemens SG, Winer JR, Sharma A, Falgàs N, Lucey BP, et al.. Wake-promoting neuromodulators in Alzheimer's disease: Implications for sleep and brain clearance. . 2026. PMID 41830117 · DOI · PMC13093618
  5. Roenneberg T, Kuehnle T, Juda M, Kantermann T, Allebrandt K, Gordijn M, et al.. Epidemiology of the human circadian clock. Sleep Med Rev. 2007. PMID 17936039 · DOI
  6. Brown RE, Basheer R, McKenna JT, Strecker RE, McCarley RW. Control of sleep and wakefulness. Physiol Rev. 2012. PMID 22811426 · DOI · PMC3621793
  7. Saper CB, Fuller PM, Pedersen NP, Lu J, Scammell TE. Sleep state switching. Neuron. 2010. PMID 21172606 · DOI · PMC3026325
  8. Lobbezoo F, Ahlberg J, Raphael KG, Wetselaar P, Glaros AG, Kato T, et al.. International consensus on the assessment of bruxism: Report of a work in progress. J Oral Rehabil. 2018. PMID 29926505 · DOI · PMC6287494
  9. Dewald JF, Meijer AM, Oort FJ, Kerkhof GA, Bögels SM. The influence of sleep quality, sleep duration and sleepiness on school performance in children and adolescents: A meta-analytic review. Sleep Med Rev. 2010. PMID 20093054 · DOI
  10. Scammell TE, Arrigoni E, Lipton JO. Neural Circuitry of Wakefulness and Sleep. Neuron. 2017. PMID 28231463 · DOI · PMC5325713

About this article

Sources
NLM Medical Subject Headings (definition, synonyms); NCBI PubMed (counts, identifiers); Europe PMC (open-access abstracts and citation counts).
Verification
Every reference was resolved against PubMed on 2026-10-31; title, year and DOI matched.
Review state
automated source verified; no human expert review has been recorded yet.
Version
1, last updated 2026-10-29.

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At a glance

22,134
PubMed records on this heading
19%
published since 2020
1,983
review articles
928
randomized trials
10
verified references below

Counts come from PubMed on 2026-10-29; they show how much has been written, not how strong the evidence is.

Related topics See all →

  • ArousalNeuroscience
  • AcromegalyNeuroscience
  • Action PotentialsNeuroscience
  • Acute PainNeuroscience
  • Alzheimer DiseaseNeuroscience

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