The DOI matches its Crossref record; 93 of the 93 references it cites were found in a registry or an index.
Sleep Spindles in Humans: Insights from Intracranial EEG and Unit Recordings
Open accessAbstract
Sleep spindles are an electroencephalographic (EEG) hallmark of non-rapid eye movement (NREM) sleep and are believed to mediate many sleep-related functions, from memory consolidation to cortical development. Spindles differ in location, frequency, and association with slow waves, but whether this heterogeneity may reflect different physiological processes and potentially serve different functional roles remains unclear. Here we used a unique opportunity to record intracranial depth EEG and single-unit activity in multiple brain regions of neurosurgical patients to better characterize spindle activity in human sleep. We find that spindles occur across multiple neocortical regions, and less frequently also in the parahippocampal gyrus and hippocampus. Most spindles are spatially restricted to specific brain regions. In addition, spindle frequency is topographically organized with a sharp transition around the supplementary motor area between fast (13-15 Hz) centroparietal spindles often occurring with slow-wave up-states, and slow (9-12 Hz) frontal spindles occurring 200 ms later on average. Spindle variability across regions may reflect the underlying thalamocortical projections. We also find that during individual spindles, frequency decreases within and between regions. In addition, deeper NREM sleep is associated with a reduction in spindle occurrence and spindle frequency. Frequency changes between regions, during individual spindles, and across sleep may reflect the same phenomenon, the underlying level of thalamocortical hyperpolarization. Finally, during spindles neuronal firing rates are not consistently modulated, although some neurons exhibit phase-locked discharges. Overall, anatomical considerations can account well for regional spindle characteristics, while variable hyperpolarization levels can explain differences in spindle frequency.
Overview
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Integrity signals
Registry
Cites
Statistics
Verification
Claims the paper makes
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
The source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“Frequencies below 20 Hz were avoided because of the potential overlap with fast sleep spindles, which can reach 18 Hz 65,66 .”
From The neural correlates of dreaming (2017) · cites this paper as [65]
Experimental — accuracy not yet measuredThe source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“Importantly, this research revealed that in the neocortex, single spindles typically express as a local phe-nomena and are restricted to specific regions and circuits, regardless of whether or not they are synchronized in phase with central thalamic spindle generation.”
From About Sleep's Role in Memory (2013) · cites this paper as [38]
Experimental — accuracy not yet measuredThe source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“As for spindles, only the classic fast spindles display the strong phase synchronization with the emergent depolarizing upstate of the slow oscillation.Slow frontal spindles, in contrast, tend to follow fast spindles by 200 -500 ms and thus occur already in the up-to-down state transition of the slow oscillation (38,826).”
From About Sleep's Role in Memory (2013) · cites this paper as [38]
Experimental — accuracy not yet measured
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
The self-check needs the paper’s full text.
Citations
Cites
(2007) · Cited by 6,530
Diekelmann, S., & Born, J. (2010) · Nature Reviews Neuroscience · doi:10.1038/nrn2762 · Cited by 3,760
Unsupervised spike detection and sorting with wavelets and superparamagnetic clustering
Quiroga, R. Q., Nadasdy, Z., & Ben-Shaul, Y. (2004) · Neural Computation · doi:10.1162/089976604774201631 · Cited by 2,119
Spike Timing–Dependent Plasticity: A Hebbian Learning Rule
Caporale, N., & Dan, Y. (2008) · Annual Review of Neuroscience · doi:10.1146/annurev.neuro.31.060407.125639 · Cited by 1,997 · cited 2×
“That spike timing is modulated by sleep spindles and is also implicated in plasticity processes (Caporale and Dan, 2008) supports the notion that spindles may have a role in synaptic plasticity (Khazipov et al., 2004).”
BackgroundHuber, R., Felice Ghilardi, M., Massimini, M., & Tononi, G. (2004) · Nature · doi:10.1038/nature02663 · Cited by 1,854
Self-citationSleep and arousal: thalamocortical mechanisms
McCormick, D. A., & Bal, T. (1997) · Annual Review of Neuroscience · doi:10.1146/annurev.neuro.20.1.185 · Cited by 1,267
The sleep slow oscillation as a traveling wave
Massimini, M., Huber, R., Ferrarelli, F., Hill, S., & Tononi, G. (2004) · The Journal of Neuroscience · doi:10.1523/jneurosci.1318-04.2004 · Cited by 1,221
Self-citationPrefrontal phase locking to hippocampal theta oscillations
Siapas, A. G., Lubenov, E. V., & Wilson, M. A. (2005) · Neuron · doi:10.1016/j.neuron.2005.02.028 · Cited by 1,014
Atlas of Electroencephalography
(1941) · JAMA · doi:10.1001/jama.1941.02820290075031 · Cited by 1,002
Gennaro, L. D., & Ferrara, M. (2003) · Sleep Medicine Reviews · doi:10.1053/smrv.2002.0252 · Cited by 957
Research works printed but not linked: 3
- Dehghani, 2010, Dynamics of excitation and inhibition, and local interactions, during slow wave sleep oscillations in rat prefrontal cortex, Soc Neurosci Abstr, 36, 44
- Jones EG (2007) The thalamus (Cambridge UP, Cambridge, UK), Ed 2.
- Steriade M (2003) Neuronal substrates of sleep and epilepsy (Cambridge UP, Cambridge, UK), Ed 1.
Cited by
Sorted by citations
Items 501–504 of 504
Pan, B., Ding, S., Geng, Y., Liang, J., Li, F., Wang, G., Li, X., Dong, Y., & Li, R. (2026) · Biosensors · doi:10.3390/bios16090467
Genuinely Blind Identification of Sleep Spindles through Trispectral Modulation AnalysisPreprint
Kovach, C. K., Gliske, S., West, L. C., Liu, J. V., Summers, M. O., Kumar, S., Gonzales, J., Cox, O., Tsang, E. W., Thompson, J. A., et al. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.08.24.746784
Sound asleep, yet still listening: auditory processing during human sleep
Oudiette, D., & Andrillon, T. (2026) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2026.09.009
Self-citationAnti-correlation time-frequency synergy and dual-granularity fusion for sleep stage classification
Pan, J., Zhao, X., Zou, X., 赵鹏军, Wang, L., & Li, C. (2026) · Frontiers in Neuroscience · doi:10.3389/fnins.2026.1944816
Metadata
- Authors
- Andrillon, T., Nir, Y., Staba, R. J., Ferrarelli, F., Cirelli, C., Tononi, G., & Fried, I.
- Affiliations
- University of Wisconsin–Madison (US)
- University of California, Los Angeles (US)
- ORCID
- Venue
- The Journal of Neuroscience
- Publisher
- Society for Neuroscience
- Published
- 2011-12-07
- Volume
- 31
- Issue
- 49
- Pages
- 17821-17834
- ISSN
- 0270-6474 (ISSN-L)
- 1529-2401
- PMC
- PMC3270580
- OpenAlex
- W2004454719
- Cited by
- 504
- References
- 93
- Citations by year
- 2026: 26 · 2025: 41 · 2024: 37 · 2023: 57 · 2022: 40 · 2021: 30 · 2020: 48 · 2019: 38 · 2018: 38 · 2017: 46 · 2016: 27 · 2015: 25 · 2014: 21 · 2013: 24 · 2012: 6
- FWCI
- 8.29
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Journal of Neuroscience (publishedVersion, cc-by-nc-sa)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Institutes of Health (DP1OD000579)
- National Institute of Neurological Disorders and Stroke (R01NS071048)
- 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 spindles, NREM sleep, single-unit activity, supplementary motor area, parahippocampal gyrus, hippocampus, spindle frequency, topographic organization
- MeSH terms
- Action Potentials, Adult, Brain, Brain Mapping, Electroencephalography, Electromyography, Electrooculography, Epilepsy, Female, Humans, Male, Middle Aged, Neurons, Sleep, Spectrum Analysis, Time Factors, Polysomnography, Nonlinear Dynamics, Young Adult, Brain Waves
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review
