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Sleep Homeostasis and Cortical Synchronization: III. A High-Density EEG Study of Sleep Slow Waves in Humans
Open accessAbstract
STUDY OBJECTIVES: The mechanisms responsible for the homeostatic decrease of slow-wave activity (SWA, defined in this study as electroencephalogram [EEG] power between 0.5 and 4.0 Hz) during sleep are unknown. In agreement with a recent hypothesis, in the first of 3 companion papers, large-scale computer simulations of the sleeping thalamocortical system showed that a decrease in cortical synaptic strength is sufficient to account for the decline in SWA. In the model, the reduction in SWA was accompanied by decreased incidence of high-amplitude slow waves, decreased wave slopes, and increased number of waves with multiple peaks. In a second companion paper in the rat, local field potential recordings during early and late sleep confirmed the predictions of the model. Here, we investigated the model's predictions in humans by using all-night high-density (hd)-EEG recordings to explore slow-wave parameters over the entire cortical mantle. DESIGN: 256-channel EEG recordings in humans over the course of an entire night's sleep. SETTING: Sound-attenuated sleep research room PATIENTS OR PARTICIPANTS: Seven healthy male subjects INTERVENTIONS: N/A. MEASUREMENTS AND RESULTS: During late sleep (non-rapid eye movement [NREM] episodes 3 and 4, toward morning), when compared with early sleep (NREM sleep episodes 1 and 2, at the beginning of the night), the analysis revealed (1) reduced SWA, (2) fewer large-amplitude slow waves, (3) decreased wave slopes, (4) more frequent multipeak waves. The decrease in slope between early and late sleep was present even when waves were directly matched by wave amplitude and slow-wave power in the background EEG. Finally, hd-EEG showed that multipeak waves have multiple cortical origins. CONCLUSIONS: In the human EEG, the decline of SWA during sleep is accompanied by changes in slow-wave parameters that were predicted by a computer model simulating a homeostatic reduction of cortical synaptic strength.
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“Independent of the decrease in amplitude, the slope of the slow waves also decreases from early to late sleep”
From About Sleep's Role in Memory (2013) · cites this paper as [994]
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“Traditionally, SWA is regarded as a marker of the homeostatically regulated sleep pressure, which increases after prolonged sleep deprivation and decreases from early to late sleep (111, 113, 994, 1269).”
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“possibly reflecting a decrease in the synchrony and speed of recruitment of neurons at the down-to-up state transition (87,360,994,1267).”
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“Specifically, the model predicted a decrease in the incidence of high-amplitude slow waves, a decrease in slope, as well as an increase in the number of multipeak waves from early to late sleep, which was con-firmed in related animal and human studies (994, 1269).”
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Research works printed but not linked: 30
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Items 201–300 of 425
Encephalopathy related to Status Epilepticus during slow Sleep: a link with sleep homeostasis?
Rubboli, G., Huber, R., Tononi, G., & Tassinari, C. A. (2019) · Epileptic Disorders · doi:10.1684/epd.2019.1059 · Cited by 24
Self-citationBonanni, E., Coscio, E. D., Maestri, M., Carnicelli, L., Tsekou, H., Economou, N. T., Paparrigopoulos, T. J., Bonakis, A., Papageorgiou, S. G., Vassilopoulos, D., et al. (2012) · Journal of Clinical Neurophysiology · doi:10.1097/wnp.0b013e318246b56d · Cited by 22
“The role of SWS delta EEG waves in memory processes has been emphasized in recent articles, which have suggested that the level of slow-wave activity might be directly proportional to the strength of cortical synapses related to cognitive processes (Riedner et al., 2007).”
BackgroundVila-Merkle, H., González-Martínez, A., Campos-Jiménez, R., Martínez‐Ricós, J., Teruel‐Martí, V., Lloret, A., Blasco‐Serra, A., & Cervera‐Ferri, A. (2023) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2023.1122163 · Cited by 29
“Slow waves are relevant to the maintenance of the general synchronization of the network, biochemical homeostasis of neurons, adaptation to environmental pressure, and memory consolidation (Maquet et al., 1997; Esser et al., 2007; Riedner et al., 2007; Neske, 2016).”
BackgroundOrigin, synchronization, and propagation of sleep slow waves in children
Castelnovo, A., Lividini, A., Riedner, B. A., Avvenuti, G., Jones, S. G., Miano, S., Tononi, G., Manconi, M., & Bernardi, G. (2023) · NeuroImage · doi:10.1016/j.neuroimage.2023.120133 · Cited by 23
“Given that slow-wave slope and globality are thought to respectively reflect short-range (Esser et al., 2007; Riedner et al., 2007; Vyazovskiy et al., 2007) and long-range (Kurth et al.”
BackgroundSelf-citationA postsleep decline in auditory evoked potential amplitude reflects sleep homeostasis
Hulse, B. K., Landsness, E. C., Sarasso, S., Ferrarelli, F., Guokas, J. J., Wanger, T., & Tononi, G. (2011) · Clinical Neurophysiology · doi:10.1016/j.clinph.2011.01.041 · Cited by 23
“Indeed, paralleling the nightly reduction in SWA, the slope of individual sleep slow waves decreases, which reflects a decline in the synchrony of up-to-down and down-to-up state transitions (Riedner et al., 2007; Vyazovskiy et al., 2009).”
Self-citationAsymmetry of sleep electrophysiological markers in patients with focal epilepsy
Sheybani, L., Mégevand, P., Roehri, N., Spinelli, L., Kleinschmidt, A. K., van Mierlo, P., Seeck, M., & Vulliémoz, S. (2023) · Brain Communications · doi:10.1093/braincomms/fcad161 · Cited by 21
Epileptic interictal discharges are more frequent during NREM slow wave downstates
Ujma, P. P., Halász, P., Kelemen, A., Fabó, D., & Erőss, L. (2017) · Neuroscience Letters · doi:10.1016/j.neulet.2017.08.020 · Cited by 21
Menicucci, D., Piarulli, A., Laurino, M., Zaccaro, A., Agrimi, J., & Gemignani, A. (2020) · Journal of Sleep Research · doi:10.1111/jsr.13117 · Cited by 23
“…fair to underline that criteria for the detection and characterization of SSOs in humans were established in the years immediately following the study of Huber and colleagues (Massimini, Huber, Ferrarelli, Hill, & Tononi, 2004; Menicucci et al., 2009; Piarulli et al., 2010; Riedner et al., 2007).”
BackgroundSousouri, G., Krugliakova, E., Skoručak, J., Leach, S., Snipes, S., Ferster, M. L., Poian, G. D., Karlen, W., & Huber, R. (2021) · SLEEP · doi:10.1093/sleep/zsab204 · Cited by 21
Self-citationAltered sleep spindles and slow waves during space shuttle missions
Koller, D. P., Kasanin, V., Flynn‐Evans, E. E., Sullivan, J. P., Dijk, D., Czeisler, C. A., & Barger, L. K. (2021) · npj Microgravity · doi:10.1038/s41526-021-00177-1 · Cited by 19
Sleep homeostasis, seizures, and cognition in children with focal epilepsy
Eriksson, M. H., Baldeweg, T., Pressler, R., Boyd, S., Huber, R., Cross, J. H., Bölsterli, B., & Chan, S. Y. S. (2022) · Developmental Medicine & Child Neurology · doi:10.1111/dmcn.15403 · Cited by 20
Self-citationBrain plasticity and sleep: Implication for movement disorders
Caverzasio, S., Amato, N., Manconi, M., Prosperetti, C., Kaelin‐Lang, A., Hutchison, W. D., & Galati, S. (2017) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2017.12.009 · Cited by 21
Mensen, A., Poryazova, R., Huber, R., & Bassetti, C. L. (2018) · Scientific Reports · doi:10.1038/s41598-018-36327-x · Cited by 22
Self-citationMensen, A., Pigorini, A., Facchin, L., Schöne, C., D’Ambrosio, S., Jendoubi, J., Jaramillo, V., Chiffi, K., Eberhard‐Moscicka, A. K., Sarasso, S., et al. (2018) · Journal of Neuroscience Methods · doi:10.1016/j.jneumeth.2018.12.011 · Cited by 22
Self-citationCan a tDCS treatment enhance subjective and objective sleep among student-athletes?
Charest, J., Marois, A., & Bastien, C. (2019) · Journal of American College Health · doi:10.1080/07448481.2019.1679152 · Cited by 20
Progressive muscle relaxation increases slow‐wave sleep during a daytime nap
Simon, K. C. N. S., McDevitt, E. A., Ragano, R., & Mednick, S. C. (2022) · Journal of Sleep Research · doi:10.1111/jsr.13574 · Cited by 24
“Considerable evidence has shown that the time spent in and the power of SWS are markers of homeostatic sleep pressure, akin to restoration needs (Borbély, 1982; Riedner et al., 2007).”
BackgroundStereotypical spatiotemporal activity patterns during slow-wave activity in the neocortex
Fucke, T., Suchanek, D., Nawrot, M. P., Seamari, Y., Heck, D. H., Aertsen, A., & Boucsein, C. (2011) · Journal of Neurophysiology · doi:10.1152/jn.00811.2010 · Cited by 19
Zhang, M., Li, R., Wang, Y., & Huang, Z. (2017) · Neural Plasticity · doi:10.1155/2017/5870735 · Cited by 19
How and when EEG reflects changes in neuronal connectivity due to time awake
Snipes, S., Meier, E., Meissner, S. N., Landolt, H., & Huber, R. (2023) · iScience · doi:10.1016/j.isci.2023.107138 · Cited by 19
“During S1, theta occupied on average 8% (1, 8) of the Fixation recording (Oddball: 10% [1, 13]; Standing: 12% [2, 16]) and more than this doubled to 22% (4, 34) during S8 (Oddball: 25% [7, 37]; Standing: 27% [8, 33]).”
BackgroundSelf-citationNovel concepts in sleep regulation
Wigren, H., & Porkka‐Heiskanen, T. (2017) · Acta Physiologica · doi:10.1111/apha.13017 · Cited by 21
“The SHY hypotheses experimentation relies on measuring synaptic strengths, as well as imaging the actual synapses during waking and sleep,(32) and the notion that the number of synapses and their strength influence the amplitude (and slope) of the slow wave activity (SWA), while neuronal synchrony is related to the nu…”
BackgroundRapid alterations in cortical protein profiles underlie spontaneous sleep and wake bouts
Vazquez, J., Hall, S. C., Witkowska, H. E., & Greco, M. A. (2008) · Journal of Cellular Biochemistry · doi:10.1002/jcb.21970 · Cited by 18
“…that following sleep deprivation (SD), there is an increase in sleep time and intensity that is proportional to the sleep lost, suggesting that a need for sleep accumulates during waking in both rodents and humans [Tobler and Borbely, 1986; Riedner et al., 2007; Vyazovskiy et al., 2007].”
BackgroundInfluentialPiarulli, A., Menicucci, D., Gemignani, A., Olcese, U., D’Ascanio, P., Pingitore, A., Bedini, R., & Landi, A. (2009) · IEEE Transactions on Biomedical Engineering · doi:10.1109/tbme.2009.2031983 · Cited by 18
Sleep Power Topography in Children with Attention Deficit Hyperactivity Disorder (ADHD)
Castelnovo, A., Lividini, A., Bernardi, G., Pezzoli, V., Foderaro, G., Ramelli, G. P., Manconi, M., & Miano, S. (2022) · Children · doi:10.3390/children9020197 · Cited by 18
“The homeostatic process is fundamental for brain plasticity [54] and is expressed by changes in the EEG power spectrum between the first and the second half of the night, mainly in the 1–9 Hz range [55], in face of a stable power topography.”
BackgroundMiano, S., Amato, N., Garbazza, C., Abbafati, M., Foderaro, G., Pezzoli, V., Ramelli, G. P., & Manconi, M. (2019) · SLEEP · doi:10.1093/sleep/zsz167 · Cited by 16
Vila-Merkle, H., González-Martínez, A., Campos-Jiménez, R., Martínez‐Ricós, J., Teruel‐Martí, V., Blasco‐Serra, A., Lloret, A., Celada, P., & Cervera‐Ferri, A. (2021) · Biomedicines · doi:10.3390/biomedicines9070783 · Cited by 16
Page, J. M., Wakschlag, L. S., & Norton, E. S. (2021) · Developmental Psychobiology · doi:10.1002/dev.22202 · Cited by 17
Examining the effects of sleep delay on depressed males and females and healthy controls
Goldschmied, J., Cheng, P., Armitage, R., & Deldin, P. J. (2014) · Journal of Sleep Research · doi:10.1111/jsr.12174 · Cited by 16
“Previous research in healthy male subjects has found that SWA is highest in the frontal regions compared to temporal and parietal regions, and have their highest amplitude during early sleep, or NREM periods 1 and 2 compared to late sleep, or NREM periods 3 and 4 (Riedner et al., 2007).”
BackgroundNatraj, N., Neylan, T. C., Yack, L. M., Metzler, T. J., Woodward, S. H., Hubachek, S. Q., Dukes, C., Udupa, N. S., Mathalon, D. H., & Richards, A. L. (2023) · Biological Psychiatry Cognitive Neuroscience and Neuroimaging · doi:10.1016/j.bpsc.2023.02.007 · Cited by 15
Somervail, R., Cataldi, J., Stephan, A., Siclari, F., & Iannetti, G. D. (2023) · SLEEP · doi:10.1093/sleep/zsad208 · Cited by 16
Clinical implication of high-density EEG sleep recordings in Parkinson’s disease
Amato, N., Caverzasio, S., & Galati, S. (2020) · Journal of Neuroscience Methods · doi:10.1016/j.jneumeth.2020.108746 · Cited by 15
Thalamic Influence on Slow Wave Slope Renormalization During Sleep
Jaramillo, V., Jendoubi, J., Maric, A., Mensen, A., Heyse, N. C., Eberhard‐Moscicka, A. K., Wiest, R., Bassetti, C. L., & Huber, R. (2021) · Annals of Neurology · doi:10.1002/ana.26217 · Cited by 14
Self-citationA role for TMS/EEG in neuropsychiatric disorders
Canali, P. (2014) · Neurology Psychiatry and Brain Research · doi:10.1016/j.npbr.2014.02.001 · Cited by 16
K-complex morphological features in male obstructive sleep apnea-hypopnea syndrome patients
Sun, L., Zhang, X., Huang, S., Liang, J., & Luo, Y. (2017) · Respiratory Physiology & Neurobiology · doi:10.1016/j.resp.2017.11.004 · Cited by 16
Oscillatory and nonoscillatory sleep electroencephalographic biomarkers of the epileptic network
Latreille, V., Corbin‐Lapointe, J., Peter‐Derex, L., Thomas, J., Lina, J., & Frauscher, B. (2024) · Epilepsia · doi:10.1111/epi.18088 · Cited by 15
Rissardo, J. P., Khalil, I., Taha, M. R., Chen, J., Sayad, R., & Caprara, A. L. F. (2025) · Medical Sciences · doi:10.3390/medsci13030113 · Cited by 19
“SWS, marked by high-voltage slow brain waves, supports memory consolidation, synaptic plasticity, and metabolic waste clearance [247].”
Januszko, P., Gmaj, B., Piotrowski, T., Kopera, M., Klimkiewicz, A., Wnorowska, A., Wołyńczyk-Gmaj, D., Brower, K. J., Wojnar, M., & Jakubczyk, A. (2020) · Drug and Alcohol Dependence · doi:10.1016/j.drugalcdep.2020.108393 · Cited by 15
Ilhan-Bayrakcı, M., Cabral‐Calderín, Y., Bergmann, T. O., Tüscher, O., & Stroh, A. (2022) · Cerebral Cortex · doi:10.1093/cercor/bhab516 · Cited by 17
“…negative signal envelope by applying a validated procedure based on the detection of negative half-waves (Mölle et al. 2002; Massimini et al. 2004; Riedner et al. 2007; Nir et al. 2011; Siclari et al. 2014; Mensen et al. 2016; Bernardi et al. 2018; Siclari et al. 2018; Avvenuti et al. 2020; Betta…”
BackgroundMethodInfluentialA region‐specific modulation of sleep slow waves on interictal epilepsy markers in focal epilepsy
Chen, C., Wang, Y., Ye, L., Xu, J., Ming, W., Liu, X., Hu, L., Ye, H., Xu, C., Wang, Y., et al. (2023) · Epilepsia · doi:10.1111/epi.17518 · Cited by 14
Targeting Slow Wave Sleep Deficiency in Late-Life Depression: A Case Series With Propofol
Rios, R. L., Kafashan, M., Hyche, O., Lenard, E. M., Lucey, B. P., Lenze, E. J., & Palanca, B. J. A. (2023) · American Journal of Geriatric Psychiatry · doi:10.1016/j.jagp.2023.03.009 · Cited by 14
Characterizing endogenous delta oscillations in human MEG
Gunasekaran, H., Azizi, L., van Wassenhove, V., & Herbst, S. K. (2023) · Scientific Reports · doi:10.1038/s41598-023-37514-1 · Cited by 13
“Delta-band activity is a prominent feature of neural dynamics traditionally observed during states of absence of consciousness, such as non-REM sleep in animals [1], [2] and humans [3]–[5].”
BackgroundThe dichotomy between low frequency and delta waves in human sleep: A reappraisal
Lanquart, J., Nardone, P., Hubain, P., Loas, G., & Linkowski, P. (2017) · Journal of Neuroscience Methods · doi:10.1016/j.jneumeth.2017.09.019 · Cited by 15
“Consistent with this observation, at the beginning of the night, when the power in the delta band is at its maximum (Preud'homme et al., 2008), the cortical neurons are tightly connected, and thus oscillate with a high degree of synchronization (Borbély et al., 1981; Riedner et al., 2007).”
BackgroundCastelnovo, A., Zago, M., Casetta, C., Zangani, C., Donati, F. L., Canevini, M., Riedner, B. A., Tononi, G., Ferrarelli, F., Sarasso, S., et al. (2020) · Schizophrenia Research · doi:10.1016/j.schres.2020.03.025 · Cited by 14
Self-citationRole of corpus callosum in sleep spindle synchronization and coupling with slow waves
Bernardi, G., Avvenuti, G., Cataldi, J., Lattanzi, S., Ricciardi, E., Polonara, G., Silvestrini, M., Siclari, F., Fabri, M., & Bellesi, M. (2021) · Brain Communications · doi:10.1093/braincomms/fcab108 · Cited by 14
“40 Hz broadband filter) and potential slow waves were identified as negative half-waves comprised between two consecutive signal zero-crossings.(3,46) Only half-waves”
BackgroundThe individual determinants of morning dream recall
Elce, V., Bergamo, D., Bontempi, G., Pedreschi, B., Bellesi, M., Handjaras, G., & Bernardi, G. (2025) · Communications Psychology · doi:10.1038/s44271-025-00191-z · Cited by 14
“Below is the model used:”
Decoding information about cognitive health from the brainwaves of sleep
Adra, N., Dummer, L., Paixão, L. G. P., Tesh, R. A., Sun, H., Ganglberger, W., Westmeijer, M., Cardoso, M. D. S., Kumar, A., Ye, E. M., et al. (2023) · Scientific Reports · doi:10.1038/s41598-023-37128-7 · Cited by 13
A probabilistic framework for a physiological representation of dynamically evolving sleep state
Dadok, V. M., Kirsch, H. E., SLEIGH, J. W., Lopour, B. A., & Szeri, A. J. (2013) · Journal of Computational Neuroscience · doi:10.1007/s10827-013-0489-x · Cited by 15
“However, the importance and meaning of this feature is highlighted in Riedner et al. (2007).”
Backgroundvan den Munckhof, B., Gefferie, S. R., van Noort, S. A. M., van Teeseling, H. C., Schijvens, M. P., Smit, W., Teunissen, N. W., Plate, J. D. J., Huiskamp, G. J. M., Leijten, F. S. S., et al. (2020) · SLEEP · doi:10.1093/sleep/zsaa088 · Cited by 14
“5–2 Hz), spikes and artifacts, based on previously published methods [19, 30].”
BackgroundMethodCserpán, D., Rosch, R. E., Biundo, S. P. L., et al. (2022) · Clinical Neurophysiology · doi:10.1016/j.clinph.2021.12.013 · Cited by 13
“14 The decrease of HFO rate with the accumulated time spent in sleep, in addition to its change with the different sleep stages, may be partly explained by the sleep-homeostatic changes in delta power showing a steady decline across the recording night (Riedner et al., 2007).”
Navarrete, M., Arthur, S., Treder, M. S., & Lewis, P. A. (2022) · NeuroImage · doi:10.1016/j.neuroimage.2022.119055 · Cited by 13
Ascent to moderate altitude impairs overnight memory improvements
Tesler, N., Latshang, T. D., Cascio, C. M. L., Stadelmann, K., Stoewhas, A., Kohler, M., Bloch, K. E., Achermann, P., & Huber, R. (2014) · Physiology & Behavior · doi:10.1016/j.physbeh.2014.11.033 · Cited by 13
“Accordingly, in humans and rats it was shown that the slope of slow waves decreases across the night [14,16, 30,31] presumably reflecting the downscaling of synaptic strength across the night.”
BackgroundMethodResultSelf-citationNeurobiology of REM Sleep, NREM Sleep Homeostasis, and Gamma Band Oscillations
McKenna, J. T., Zielinski, M. R., & McCarley, R. W. (2017) · Springer eBooks · doi:10.1007/978-1-4939-6578-6_5 · Cited by 15
Modelling Sleep and General Anaesthesia
SLEIGH, J. W., Voss, L. J., Steyn‐Ross, M. L., Steyn‐Ross, D. A., & Wilson, M. T. (2011) · Springer eBooks · doi:10.1007/978-1-4614-0173-5_2 · Cited by 15
“…to such an extent that the balance shifts in favor of the aminergic activating systems; which then inhibit the VLPO and initiate a positive feedback of arousal that is the inverse of that described above when the person falls asleep (Rempe et al. 2010; Riedner et al. 2007; Wilson et al. 2005).”
BackgroundBeta spectral power during sleep is associated with impaired recall of extinguished fear
Denis, D., Bottary, R. M., Cunningham, T. J., Drummond, S. P. A., & Straus, L. D. (2023) · SLEEP · doi:10.1093/sleep/zsad209 · Cited by 12
“5–4 Hz PSD) has been found to be a reliable marker of homeostatic sleep pressure and is elevated in recovery sleep following sleep deprivation [42].”
BackgroundCarvalho, D. Z., Křemen, V., Mívalt, F., Louis, E. K. S., McCarter, S. J., Bukartyk, J., Przybelski, S. A., Kamykowski, M. G., Spychalla, A. J., Machulda, M. M., et al. (2024) · Brain Communications · doi:10.1093/braincomms/fcae354 · Cited by 12
“This methodological difference is relevant because homeostatic pressure for sleep is mostly reflected in SWA, 64-69 with higher contribution, amplitudes and slopes at the beginning of sleep, 39,70 with decay during the course of sleep.”
BackgroundMethodResultInfluentialBlindfolding during wakefulness causes decrease in sleep slow wave activity
Korf, E. M., Mölle, M., Jan Born, A., & Ngo, H.-V. V. (2017) · Physiological Reports · doi:10.14814/phy2.13239 · Cited by 11
“This hypothesis is supported not only by a number of computational modeling studies (Esser et al. 2007; Riedner et al. 2007; Olcese et al. 2010), but also by experimental data showing correlations between SWA and changes in cortical synaptic density across the lifespan.”
ResultComputational models of memory consolidation and long-term synaptic plasticity during sleep
Rennó‐Costa, C., da Silva, A. C. C., Blanco, W., & Ribeiro, S. T. G. (2018) · Neurobiology of Learning and Memory · doi:10.1016/j.nlm.2018.10.003 · Cited by 12
“…and network mechanisms underlying slow-wave activity observed during sleep and the transition to an awake brain state (Bazhenov, Timofeev, Steriade, & Sejnowski, 2002; Compte, SanchezVives, McCormick, & Wang, 2003; Esser, Hill, & Tononi, 2007, 2009; Hill & Tononi, 2005; Komarov et al., 2018).”
Parapatics, S., Anderer, P., Gruber, G., Saletu, B., Saletu-Zyhlarz, G. M., & Dorffner, G. (2015) · Somnologie - Schlafforschung und Schlafmedizin · doi:10.1007/s11818-015-0701-5 · Cited by 11
Development of slow oscillation–spindle coupling from infancy to toddlerhood
Kurz, E., Bastian, L., Mölle, M., Jan Born, A., & Friedrich, M. (2024) · SLEEP Advances · doi:10.1093/sleepadvances/zpae084 · Cited by 14
“The slope of the SO primarily reflects the extent of synchronization in the underlying neuronal membrane potentials and has been proposed as an indicator of synaptic strength [44, 45].”
Local Patterns of Sleep and Wakefulness
Bernardi, G., & Siclari, F. (2019) · Handbook of behavioral neuroscience · doi:10.1016/b978-0-12-813743-7.00003-7 · Cited by 14
Navarrete, M., Osorio-Forero, A., Gómez, A., Henao, D., Segura-Quijano, F. E., Quyen, M. L. V., & Valderrama, M. (2023) · SLEEP · doi:10.1093/sleep/zsad110 · Cited by 13
Miano, S., & Datta, A. (2019) · European Journal of Pediatrics · doi:10.1007/s00431-019-03413-9 · Cited by 12
Cucchiara, F., Frumento, P., Banfi, T., Sesso, G., Galante, M. D., D’Ascanio, P., Valvo, G., Sicca, F., & Faraguna, U. (2019) · SLEEP · doi:10.1093/sleep/zsz255 · Cited by 11
“Thus, the period of slow waves, which is generally modulated by extracellular factors such as the persistent Na+ current and the depolarization-activated K+ current [33], could be reliably detected in ASD/AEP children carrying GoF Kir4.”
BackgroundInfluentialGefferie, S. R., Maric, A., Critelli, H., Guéden, S., Kurlemann, G., Kurth, S., Nosadini, M., Plecko, B., Ringli, M., Rostasy, K. M., et al. (2021) · NeuroImage · doi:10.1016/j.neuroimage.2021.118281 · Cited by 10
Self-citationImpairment of sleep homeostasis in cervical dystonia patients
Caverzasio, S., Amato, N., Chiaro, G., Staedler, C., Kaelin‐Lang, A., & Galati, S. (2022) · Scientific Reports · doi:10.1038/s41598-022-10802-y · Cited by 12
Propagating population activity patterns during spontaneous slow waves in the thalamus of rodents
Horváth, C., Ulbert, I., & Fiáth, R. (2023) · NeuroImage · doi:10.1016/j.neuroimage.2023.120484 · Cited by 10
Stephan, A., Cataldi, J., Virk, A. S., & Siclari, F. (2025) · Current Biology · doi:10.1016/j.cub.2025.06.064 · Cited by 11
Focal limbic sources create the large slow oscillations of the EEG in human deep sleep
Morgan, K. K., Hathaway, E., Carson, M., Fernández-Corazza, M., Shusterman, R., Luu, P., & Tucker, D. M. (2021) · Sleep Medicine · doi:10.1016/j.sleep.2021.07.028 · Cited by 10
Darchia, N., Campbell, I. G., Basishvili, T., Eliozishvili, M., Tchintcharauli, T., Oniani, N., Sakhelashvili, I., & Feinberg, I. (2022) · SLEEP · doi:10.1093/sleep/zsac163 · Cited by 9
Lafrenière, A., Lina, J., Hernandez, J., Bouchard, M., Gosselin, N., & Carrier, J. (2022) · SLEEP · doi:10.1093/sleep/zsac246 · Cited by 12
Oz, O., Matityahu, L., Mizrahi-Kliger, A. D., Kaplan, A., Berkowitz, N., Tiroshi, L., Bergman, H., & Goldberg, J. A. (2022) · eLife · doi:10.7554/elife.76039 · Cited by 9
Picot, A., Whitmore, H. R., & Chapotot, F. (2012) · IEEE Transactions on Biomedical Engineering · doi:10.1109/tbme.2012.2210894 · Cited by 10
“explicit limit on the lower SSW frequency, our results may differ from those of other variants without full-wave duration criteria, such as in [15] and in [16].”
ResultShorter duration of non‐rapid eye movement sleep slow waves inEphA4knockout mice
Freyburger, M., Poirier, G., Carrier, J., & Mongrain, V. (2017) · Journal of Sleep Research · doi:10.1111/jsr.12532 · Cited by 10
“Precisely, the number of SW per minute (i.e. density), SW amplitude and slope between the positive and negative phases are increased after wakefulness compared with sleep (Bersagliere and Achermann, 2010; Riedner et al., 2007; Vyazovskiy et al., 2009).”
BackgroundInfraslow coordination of slow wave activity through altered neuronal synchrony
Dash, M. B. (2019) · SLEEP · doi:10.1093/sleep/zsz170 · Cited by 9
“Propagation failures can result in the absence of slow waves within a local cortical circuit despite the presence of a slow wave elsewhere [7, 8], while the convergence of independently generated local slow waves may give rise to multipeak waves [9, 10].”
BackgroundResultSquarcio, F., Tononi, G., & Cirelli, C. (2024) · European Journal of Neuroscience · doi:10.1111/ejn.16460 · Cited by 11
“…synaptic weakening linked to the ON/OFF pattern of the slow waves would have the appealing feature of being self-limiting (Olcese et al., 2010), like the slow waves themselves that are large and global in early sleep but local and small in late sleep (Riedner et al., 2007; Vyazovskiy et al., 2007).”
Self-citationMarsh, B. M., Navas-Zuloaga, M. G., Rosen, B. Q., Sokolov, Y., Delanois, J. E., Gonzalez, O. C., Krishnan, G. P., Halgren, E., & Bazhenov, M. (2024) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1012245 · Cited by 10
“Furthermore, [53] and [52] have reported that the occurrence of multi-peak slow waves, where each peak has a different origin, increases over the course of sleep, which was proposed to result from a gradual net decrease in the strength of cortico-cortical connections [28] (however, see [75]).”
BackgroundHemispherotomy leads to persistent sleep-like slow waves in the isolated cortex of awake humans
Colombo, M. A., Favaro, J., Mikulan, E., Pigorini, A., Zauli, F. M., Sartori, I., d’Orio, P., Castana, L., Toldo, I., Sartori, S., et al. (2025) · PLoS Biology · doi:10.1371/journal.pbio.3003060 · Cited by 10
“Different from mastoid-referenced EEG, where only half-period fluctuations within the negative polarity are considered [88–90], in our case of bipolar channels we considered full-period signal fluctuations across positive and negative polarity.”
Self-citationMenicucci, D., Piarulli, A., Allegrini, P., Bedini, R., Bergamasco, M., Laurino, M., Sebastiani, L., & Gemignani, A. (2015) · International Journal of Psychophysiology · doi:10.1016/j.ijpsycho.2015.05.006 · Cited by 9
“Herein we choose to focalize on the first sleep cycle because the SSO are homeostatically regulated and thus maximally expressed during the first sleep cycle (Riedner et al., 2007).”
InfluentialGalati, S., Song, W., Orbán, G., Luft, A. R., & Kaelin‐Lang, A. (2017) · Experimental Neurology · doi:10.1016/j.expneurol.2017.12.004 · Cited by 9
Ren, G., Yan, J., Sun, Y., et al. (2020) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2020.00335 · Cited by 9
Saadatmehr, B., Edalati, M., Routier, L., Mahmoudzadeh, M., Safaie, J., Kongolo, G., Ghostine, G., Wallois, F., & Moghimi, S. (2022) · Cerebral Cortex · doi:10.1093/cercor/bhac067 · Cited by 10
“S1) were determined as the ratio between the absolute value of the SW trough, corresponding to the peak of the negative SW half-wave, and the time interval from the previous and to the next zero crossing (Riedner et al. 2007; Dehnavi et al. 2021).”
BackgroundRay, L. B., Baena, D., & Fogel, S. M. (2024) · Journal of Neuroscience Methods · doi:10.1016/j.jneumeth.2024.110162 · Cited by 8
Guay, C. S., Labonte, A. K., Montana, M. C., Landsness, E. C., Lucey, B. P., Kafashan, M., Haroutounian, S., Avidan, M. S., Brown, E. N., & Palanca, B. J. A. (2021) · Nature and Science of Sleep · doi:10.2147/nss.s293160 · Cited by 8
Bergamo, D., Handjaras, G., Petruso, F., Talami, F., Ricciardi, E., Benuzzi, F., Vaudano, A. E., Meletti, S., Bernardi, G., & Betta, M. (2023) · Sleep Medicine · doi:10.1016/j.sleep.2023.12.001 · Cited by 10
Goldbart, A. D., Arazi, A., Golan‐Tripto, I., Levinsky, Y., Scheuerman, O., & Tarasiuk, A. (2020) · Journal of Clinical Sleep Medicine · doi:10.5664/jcsm.8678 · Cited by 8
“5,8,14 It is established that decreased sleep pressure can generate deeper SWS, which can be quanti fi ed by the power of SWA, 5 and reduced sleep pressure may lead to longer sleep latency.”
Local sleep: A new concept in brain plasticity
Avvenuti, G., & Bernardi, G. (2022) · Handbook of clinical neurology · doi:10.1016/b978-0-12-819410-2.00003-5 · Cited by 10
Increased cortical involvement and synchronization during CAP A1 slow waves
Ujma, P. P., Halász, P., Simor, P., Fabó, D., & Ferri, R. (2018) · Brain Structure and Function · doi:10.1007/s00429-018-1703-4 · Cited by 9
BOLD signatures of sleepPreprint
Song, C., Boly, M., Tagliazucchi, E., Laufs, H., & Tononi, G. (2019) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/531186 · Cited by 8
“To test this hypothesis, we detected individual occurrences of spindles and slow waves using methods described in (Ferrarelli et al., 2007, Riedner et al., 2007).”
BackgroundMethodSelf-citationVaquer-Alicea, A. D. C., Yu, J., Liu, H., & Lucey, B. P. (2023) · SLEEP Advances · doi:10.1093/sleepadvances/zpad047 · Cited by 7
Cortical hierarchy underlying homeostatic sleep pressure alleviation
Zou, Q., Zou, G., Wang, S., Wang, Y., Xu, J., Long, Y., Zhou, S., Wu, X., Yang, G., Qin, L., et al. (2025) · Nature Communications · doi:10.1038/s41467-025-64989-5 · Cited by 10
Aerobic fitness and the sleeping brain of adolescents—a pilot study
Neikrug, A. B., Mander, B. A., Radom‐Aizik, S., Chen, I. Y., Stehli, A., Lui, K. K., Chappel-Farley, M. G., Dave, A., & Benca, R. M. (2021) · SLEEP Advances · doi:10.1093/sleepadvances/zpab005 · Cited by 8
“Data preprocessing, including semi-automated artifact rejection [33, 35, 36], and analyses of sleep data focusing on NREM stages (N2 and N3) were performed using custom MATLAB scripts as previously described [37].”
MethodSleep Slow Oscillations and Cortical Maturation
Kurth, S., & Huber, R. (2012) · Elsevier eBooks · doi:10.1016/b978-0-12-384995-3.00010-1 · Cited by 8
Self-citationDetection of characteristic waves of sleep EEG by continuous wavelet transform
Koley, B. L., & Dey, D. (2012) · National Conference on Computing and Communication Systems (NCCCS) · doi:10.1109/ncccs.2012.6413025 · Cited by 7
Ng, T.-P., Noh, E., & Spencer, R. M. C. (2024) · eLife · doi:10.7554/elife.101992 · Cited by 16
Sleep and wake in a model of the thalamocortical system with Martinotti cells
Bugnon, T., Mayner, W. G. P., Cirelli, C., & Tononi, G. (2024) · European Journal of Neuroscience · doi:10.1111/ejn.15836 · Cited by 6
“Following prolonged wake, local SWs can appear (Vyazovskiy et al., 2011), and SWA decreases throughout sleep, with a reduction in the slope and the frequency of occurrence of large waves (Riedner et al., 2007; Vyazovskiy et al., 2007).”
BackgroundMethodSelf-citationLeduc, T., Alami, H. E., Bougadir, K., Bélanger-Nelson, E., & Mongrain, V. (2024) · Molecular Autism · doi:10.1186/s13229-024-00594-5 · Cited by 6
“SW properties such as density, amplitude, and slope have been reported to be higher under elevated homeostatic sleep pressure in both humans and rodents [45, 56–60].”
BackgroundYang, F. N., Picchioni, D., de Zwart, J. A., Wang, Y., van Gelderen, P., & Duyn, J. H. (2024) · eLife · doi:10.7554/elife.98739 · Cited by 6
“ICA cleaning and slow wave auto-detection script were applied to EEG signals (Betta et al., 2021; Mensen et al., 2016; Riedner et al., 2007).”
MethodLi, Z., Wang, J., Tang, C., Wang, P., Ren, P., Li, S., Yi, L., Liu, Q., Sun, L., Li, K. X., et al. (2024) · Communications Biology · doi:10.1038/s42003-024-06941-9 · Cited by 8
“As synaptic downscaling occurs during sleep, there is a corresponding reduction in the slow wave slope, mirroring the decrease in synaptic strength and density 51 .”
Plante, D. T., Goldstein, M. R., Cook, J. D., Smith, R., Riedner, B. A., Rumble, M. E., Jelenchick, L. A., Roth, A., Tononi, G., Benca, R. M., et al. (2016) · International Journal of Psychophysiology · doi:10.1016/j.ijpsycho.2016.01.003 · Cited by 6
“Slow wave detection was performed using similar methods previously described (Riedner et al., 2007).”
BackgroundMethodResultInfluentialSelf-citationChylinski, D., Narbutas, J., Balteau, E., Collette, F., Bastin, C., Berthomier, C., Salmon, É., Maquet, P., Carrier, J., Phillips, C. L. M., et al. (2022) · SLEEP · doi:10.1093/sleep/zsac178 · Cited by 6
NREM Sleep Regulation From Neuronal Assembly to Ion
Gompf, H. S., & Anaclet, C. (2019) · Handbook of behavioral neuroscience · doi:10.1016/b978-0-12-813743-7.00009-8 · Cited by 6
Metadata
- Authors
- Riedner, B. A., Vyazovskiy, V. V., Huber, R., Massimini, M., Esser, S., Murphy, M., & Tononi, G.
- Affiliations
- University of Wisconsin–Madison (US)
- Corresponding author
- Tononi, Giulio
- ORCID
- Venue
- Sleep
- Publisher
- Oxford University Press (OUP)
- Published
- 2007-12
- Volume
- 30
- Issue
- 12
- Pages
- 1643-1657
- ISSN
- 0161-8105 (ISSN-L)
- 1550-9109
- PMC
- PMC2276133
- OpenAlex
- W1577987636
- Cited by
- 424
- References
- 0
- Citations by year
- 2026: 25 · 2025: 27 · 2024: 28 · 2023: 24 · 2022: 31 · 2021: 31 · 2020: 24 · 2019: 28 · 2018: 20 · 2017: 28 · 2016: 16 · 2015: 20 · 2014: 28 · 2013: 24 · 2012: 19
- FWCI
- 5.00
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- SLEEP (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Open Research (University of Surrey) (submittedVersion)
- Surrey Research Insight Open Access (The University of Surrey) (submittedVersion)
- Funders
- Schweizerischer Nationalfonds zur Förderung der Wissenschaftlichen Forschung (106264)
- National Institutes of Health
- National Institute of General Medical Sciences (T32GM007507)
- National Institute of Diabetes and Digestive and Kidney Diseases (T90DK070079)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Sleep and Wakefulness Research
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Social Sciences › Psychology › Experimental and Cognitive Psychology › Sleep and related disorders
- Keywords
- sleep slow waves, slow-wave activity, sleep homeostasis, cortical synchronization, thalamocortical system, synaptic efficacy, NREM sleep, high-density EEG
- MeSH terms
- Adult, Brain Mapping, Cerebral Cortex, Circadian Rhythm, Computer Simulation, Cortical Synchronization, Electroencephalography, Evoked Potentials, Fourier Analysis, Homeostasis, Humans, Male, Nerve Net, Reference Values, Signal Processing, Computer-Assisted, Sleep, Sleep, REM, Synapses, Thalamus
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