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Widespread Changes in Synaptic Markers as a Function of Sleep and Wakefulness in Drosophila
Open accessAbstract
Sleep is universal, strictly regulated, and necessary for cognition. Why this is so remains a mystery, although recent work suggests that sleep, memory, and plasticity are linked. However, little is known about how wakefulness and sleep affect synapses. Using Western blots and confocal microscopy in Drosophila , we found that protein levels of key components of central synapses were high after waking and low after sleep. These changes were related to behavioral state rather than time of day and occurred in all major areas of the Drosophila brain. The decrease of synaptic markers during sleep was progressive, and sleep was necessary for their decline. Thus, sleep may be involved in maintaining synaptic homeostasis altered by waking activities.
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“The outcome that better “content-relevant sleep” leads to improved performance is supported by previous controlled studies on the role of sleep in memory consolidation.14,15”
From Sleep quality, duration, and consistency are associated with better academic performance in college students (2019) · cites this paper as [14]
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“the general understanding is that specific synaptic connections that were active during awake-periods are strengthened during sleep, allowing for the consolidation of memory, and synaptic connections that were inactive are weakened.5,14,15”
From Sleep quality, duration, and consistency are associated with better academic performance in college students (2019) · cites this paper as [14]
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“Suppressing expression of LTP-related genes in rats by chronic lesion of the noradrenergic system produced a strong reduction in the homeostatic SWA response (see, e.g., Refs.205 and 457, for similar results in flies).”
From About Sleep's Role in Memory (2013) · cites this paper as [457]
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“Sleep is thought to play a crucial and specific role in memory consolidation.”
From Sleep quality, duration, and consistency are associated with better academic performance in college students (2019) · cites this paper as [14]
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“The answer that emerges from the above arguments is that sleep is an optimization process that is disclosed by the nightly removal of sensory perturbations; in other words, the brain can take itself off-line with impunity, so that synaptic plasticity and homoeostasis (Gilestro et al., 2009) can reduce the complexity it has accrued during wakefulness.”
From Virtual reality and consciousness inference in dreaming (2014) · cites this paper as Gilestro et al., 2009
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“Thus, mechanisms of synaptic homeostasis [62] and plasticity to maintain the number of synapses in check (including the decrease of synaptic markers during sleep [63]), as well as sparse coding, with only a small proportion of neurons firing at high frequencies at any moment [49], [64], [65], may be direct consequences to brain function of a limiting and size-invariant, fixed energy budget per”
From Scaling of Brain Metabolism with a Fixed Energy Budget per Neuron: Implications for Neuronal Activity, Plasticity and Evolution (2011) · cites this paper as [63]
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Sleep function and synaptic homeostasis
Tononi, G., & Cirelli, C. (2006) · Sleep Medicine Reviews · doi:10.1016/j.smrv.2005.05.002 · Cited by 2,151
Self-citationHuber, R., Felice Ghilardi, M., Massimini, M., & Tononi, G. (2004) · Nature · doi:10.1038/nature02663 · Cited by 1,854
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El-Husseini, A. E.-D., Schnell, E., Chetkovich, D. M., Nicoll, R. A., & Bredt, D. S. (2000) · Science · doi:10.1126/science.290.5495.1364 · Cited by 1,303
Correlates of Sleep and Waking in Drosophila melanogaster
Shaw, P. J., Cirelli, C., Greenspan, R. J., & Tononi, G. (2000) · Science · doi:10.1126/science.287.5459.1834 · Cited by 1,300
Self-citationRest in Drosophila is a sleep-like state
Hendricks, J. C., Finn, S. M., Panckeri, K. A., Chavkin, J., Williams, J. A., Sehgal, A., & Pack, A. I. (2000) · Neuron · doi:10.1016/s0896-6273(00)80877-6 · Cited by 1,050
Wagh, D. A., Rasse, T. M., Asan, E., Hofbauer, A., Schwenkert, I., Dürrbeck, H., Buchner, S., Dabauvalle, M.-C., Schmidt, M., Qin, G., et al. (2006) · Neuron · doi:10.1016/j.neuron.2006.02.008 · Cited by 908
Woods, D. F., & Bryant, P. J. (1991) · Cell · doi:10.1016/0092-8674(81)90009-x · Cited by 898
Vyazovskiy, V. V., Cirelli, C., Pfister-Genskow, M., Faraguna, U., & Tononi, G. (2008) · Nature Neuroscience · doi:10.1038/nn2035 · Cited by 837
Self-citationCirelli, C., & Tononi, G. (2008) · PLoS Biology · doi:10.1371/journal.pbio.0060216 · Cited by 646
Self-citationEhrlich, I., & Malinow, R. (2004) · The Journal of Neuroscience · doi:10.1523/jneurosci.4733-03.2004 · Cited by 567
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Additional file 2 of Time of day as a critical variable in biology
Nelson, R. J., Bumgarner, J. R., Liu, J. A., Love, J. A., Meléndez‐Fernández, O. H., Becker‐Krail, D. D., Walker, W. H., Walton, J. C., DeVries, A. C., & Prendergast, B. J. (2022) · Figshare · doi:10.6084/m9.figshare.20076777
Wu, H., Maekawa, Y., Eno, S., Kondo, S., Yamagata, N., & Tanimoto, H. (2026) · eLife · doi:10.7554/elife.107663.2
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Huang, S., Fuscà, D., Moron, V., Escher, M., Ramesh, N., Toppe, D., Zhao, Z., Turrel, O., Wu, Y., Liu, S., et al. (2026) · Current Biology · doi:10.1016/j.cub.2026.09.007
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Metadata
- Authors
- Gilestro, G. F., Tononi, G., & Cirelli, C.
- Affiliations
- University of Wisconsin–Madison (US)
- Corresponding author
- Tononi, Giulio
- Cirelli, Chiara
- Venue
- Science
- Published
- 2009-04-03
- Volume
- 324
- Issue
- 5923
- Pages
- 109-112
- ISSN
- 0036-8075 (ISSN-L)
- 1095-9203
- PMC
- PMC2715914
- OpenAlex
- W2015113576
- Cited by
- 360
- References
- 29
- Citations by year
- 2026: 14 · 2025: 23 · 2024: 16 · 2023: 14 · 2022: 17 · 2021: 8 · 2020: 15 · 2019: 20 · 2018: 11 · 2017: 22 · 2016: 19 · 2015: 18 · 2014: 22 · 2013: 24 · 2012: 26
- FWCI
- 18.55
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Science (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- Funders
- National Institutes of Health (DP1OD000579)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Sleep and Wakefulness Research
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurobiology and Insect Physiology Research
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Keywords
- sleep, wakefulness, synaptic markers, synaptic homeostasis, Drosophila, behavioral state, memory consolidation, neuroplasticity
- MeSH terms
- Animals, Biological Clocks, Brain, Circadian Rhythm, Drosophila melanogaster, Female, Homeostasis, Male, Sleep, Sleep Deprivation, Synapses, Wakefulness, Blotting, Western, Synapsins, Microscopy, Confocal, Models, Animal, Tumor Suppressor Proteins, Drosophila Proteins, Qa-SNARE Proteins
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