Sleep—A brain-state serving systems memory consolidation

In the paper

“Spontaneous memory reactivations can be decoded from the EEG signal during SO-spindle complexes after the participants had learned word-image pairs using multivariate classification 114 (Figure 3).”

What the sources say

• Disputed

  The passages describing the methods and learning task state that this study recorded cortical and hippocampal activity in rats trained on a spatial cheeseboard maze, not EEG from participants learning word–image pairs. The reported result was stronger hippocampal–cortical ripple coupling during postlearning sleep; the source does not report decoding memory reactivations with multivariate classification. (LLM, conf=0.86)

  Checked by a reasoning model — may be wrong.

  cites [114] 114. Khodagholy, D., Gelinas, J.N., and Buzsa ´ki, G. (2017). Learning- enhanced coupling between ripple oscillations in association cortices and hippocampus. Science 358, 369-372. 192. sourceExperimental — accuracy not yet measured

In the paper

“Subsequently, slow oscillatory activity shows an inverted U-shaped trajectory peaking during early adolescence”

What the sources say

• Disputed

  The Introduction (p. 3) says slow wave activity reaches its maximum during childhood and declines throughout adolescence, contrary to a peak in early adolescence. The Age groups section (p. 5) places the maximum in the 5–8.9-year age group. (LLM, conf=0.88)

  Checked by a reasoning model — may be wrong.

  cites [253] sourceExperimental — accuracy not yet measured

In the paper

“Rat pups showed increases in spindle density in parallel with the emergence of hippocampal object-in-place memories during early development.275”

What the sources say

• Disputed

  The Results (Postnatal Hippocampal-Dependent Memory Onset; page unavailable) place the emergence of object-place recognition memory at P32. But the Discussion (page unavailable) explicitly reports no reliable developmental change in spindle density; it relates higher spindle density to better memory performance instead. That contradicts the claimed developmental increase in spindle density. (LLM, conf=0.89)

  Checked by a reasoning model — may be wrong.

  cites [275] sourceExperimental — accuracy not yet measured

In the paper

“The memory-related enhancement of hippocampal activation during sleep was still present when tested 1 week after the encoding session.”

What the sources say

• Partly supported

  The Results (fMRI results; page unavailable) report an association between temporal memory and hippocampal activation during sleep. The Methods (Procedure; page unavailable) place imaging after the one-week memory test, but also describe renewed exposure to the game and song before imaging. The paper therefore does not establish that an enhancement measured at encoding persisted unchanged for one week. (LLM, conf=0.86)

  cites [272] sourceExperimental — accuracy not yet measured
• Disputed

  The Results section states that left hippocampal activation for newly learned words was associated with puppet-name memory only at the same-session test. At the one-week memory test, that association was not statistically significant (rs(30) = .33, p = .07). The paper therefore does not report that a memory-related hippocampal effect remained present at one week. (LLM, conf=0.90)

  Checked by a reasoning model — may be wrong.

  cites [273] sourceExperimental — accuracy not yet measured

In the paper

“a number of studies revealed signs of an improved abstraction of rules and regularities at a later test session when participants had slept after encoding of the stimulus materials, in comparison with a wake control condition.”

What the sources say

• ?Not checked — no full text of the sourcecites [35] 35. Wagner, U., Gais, S., Haider, H., Verleger, R., and Born, J. (2004). Sleep in- spires insight. Nature 427, 352-355. https://doi.org/10.1038/nature02223. source
• Supported

  The Discussion (page not provided) says that prior research found sleep supports abstraction and reports this study’s own sleep-dependent temporal abstraction of sequence knowledge. It states that prediction strength differed significantly after sleep compared with wakefulness at the later test. (LLM, conf=0.90)

  cites [36] 36. Lutz, N.D., Wolf, I., H€ ubner, S., and Rauss, K. (2018). Sleep strengthens predictive sequence coding. J. Neurosci. 17, 8989-9000. https://doi.org/ 10.1523/JNEUROSCI.1352-18.2018. source
• Partly supported

  The Abstract (page not provided) reports that numerous studies found better memory after sleep than after comparable time awake, and describes one study in which sleepers were more likely than awake participants to discover a hidden shortcut. It does not establish that a number of studies specifically found improved abstraction of rules and regularities at a later test; this paper’s own comparison is between cued and uncued puzzles among sleepers. (LLM, conf=0.83)

  cites [37] 37. Sanders, K.E.G., Osburn, S., Paller, K.A., and Beeman, M. (2019). Targeted memory reactivation during sleep improves next-day problem solving. Psychol. Sci. 30, 1616-1624. https://doi.org/10.1177/0956797619873344. sourceExperimental — accuracy not yet measured

In the paper

“sleep-dependent gains in memories originally classified as ''non-hippocampus dependent'' can be abolished by suppressing hippocampal function during sleep.”

What the sources say

• ?Not checked — no full text of the sourcecites [66] 66. Sawangjit, A., Oyanedel, C.N., Niethard, N., Salazar, C., Born, J., and In- ostroza, M. (2018). The hippocampus is crucial for forming non-hippo- campal long-term memory during sleep. Nature 564, 109-113. https:// doi.org/10.1038/s41586-018-0716-8. source
• Partly supported

  The Discussion reports that patients with hippocampal damage learned the motor sequence task initially but, unlike controls, did not improve overnight. It also says the 24-hour design cannot establish whether the effect was specifically sleep-dependent. The study did not suppress hippocampal function during sleep; it compared patients with existing damage to controls. (LLM, conf=0.88)

  cites [74] 74. Schapiro, A.C., Reid, A.G., Morgan, A., Manoach, D.S., Verfaellie, M., and Stickgold, R. (2019). The hippocampus is necessary for the consol- idation of a task that does not require the hippocampus for initial learning. Hippocampus 29, 1091-1100. https://doi.org/10.1002/hipo.23101. sourceExperimental — accuracy not yet measured

In the paper

“with sleep exerting a more gradual ameliorating effect on the affective response.22,23”

What the sources say

• ?Not checked — no full text of the sourcecites [22] 22. Zeng, S., Lin, X., Wang, J., and Hu, X. (2021a). Sleep's short-term mem- ory preservation and long-term affect depotentiation effect in emotional memory consolidation: behavioral and EEG evidence. Sleep 44, zsab155. https://doi.org/10.1093/sleep/zsab155. source
• Partly supported

  The Sleep post-retrieval/during reconsolidation section reports a reduced heart-rate response after a nap, supporting an ameliorating effect on responses to an aversive memory. But the stimulus was an autobiographical memory script, not aversive pictures, and the effect did not persist after one week; the passage does not establish a gradual effect. (LLM, conf=0.83)

  cites [23] 23. Cunningham, T.J., Stickgold, R., and Kensinger, E.A. (2022). Investi- gating the effects of sleep and sleep loss on the different stages of episodic emotional memory: A narrative review and guide to the future. Front. Behav. Neurosci. 16, 910317. https://doi.org/10.3389/fnbeh. 2022.910317. sourceExperimental — accuracy not yet measured

In the paper

“tasks requiring motor adaptation (such as pursuit rotor task) might represent an exception as they do not seem to benefit from sleep.”

What the sources say

• Partly supported

  The Discussion reports no significant sleep effect on consolidation of the force-field motor adaptation task. The Introduction notes that some visuomotor adaptation tasks do benefit from sleep, and the paper does not study the pursuit rotor task. It therefore supports the possibility of an exception, but not the broader implication about motor adaptation tasks or the named example. (LLM, conf=0.85)

  cites [19] 19. Th€ urer, B., Weber, F.D., Born, J., and Stein, T. (2018). Variable training but not sleep improves consolidation of motor adaptation. Sci. Rep. 8, 15977. https://doi.org/10.1038/s41598-018-34225-w. sourceExperimental — accuracy not yet measured

In the paper

“Studies experimentally enhancing or suppressing SO activity through closed-loop auditory or electrical stimulation support the view of a causal role of SOs in enhancing the consolidation process.”

What the sources say

• ?Not checked — no full text of the sourcecites [50] 50. Marshall, L., Helgado ´ttir, H., Mo ¨lle, M., and Born, J. (2006). Boosting slow oscillations during sleep potentiates memory. Nature 444, 610-613. https://doi.org/10.1038/nature05278. source
• ?Not checked — no full text of the sourcecites [51] 51. Ngo, H.V., Martinetz, T., Born, J., and Mo ¨lle, M. (2013). Auditory closed- loop stimulation of the sleep slow oscillation enhances memory. Neuron 78, 545-553. https://doi.org/10.1016/j.neuron.2013.03.006. source
• Partly supported

  Box 1 reports that phase-locked auditory stimulation enhanced slow oscillations and improved overnight memory retention, supporting a causal role in consolidation. The Introduction reports improved retention after electrical induction of slow oscillations, but says those electrical studies were not synchronized with endogenous activity. The cited passages do not establish memory benefits from suppressing slow oscillations or from closed-loop electrical stimulation. (LLM, conf=0.88)

  cites [52] 52. Harrington, M.O., and Cairney, S.A. (2021). Sounding it out: auditory stimulation and overnight memory processing. Curr. Sleep Med. Rep. 7, 112-119. https://doi.org/10.1007/s40675-021-00207-0. sourceExperimental — accuracy not yet measured

In the paper

“contributions of REM sleep appear to be particularly strong for the consolidation of emotional materials.”

What the sources say

• Supported

  The Discussion reports that the emotional-memory benefit was associated with REM-sleep characteristics and was selective to REM, with no comparable association for other sleep stages. This supports the hedged claim that REM sleep contributes particularly strongly to consolidating emotional material. (LLM, conf=0.93)

  cites [25] 25. Nishida, M., Pearsall, J., Buckner, R.L., and Walker, M.P. (2009). REM sleep, prefrontal theta, and the consolidation of human emotional memory. Cereb. Cortex 19, 1158-1166. https://doi.org/10.1093/cercor/bhn155. source
• ?Not checked — no full text of the sourcecites [43] 43. Groch, S., Zinke, K., Wilhelm, I., and Born, J. (2015). Dissociating the con- tributions of slow-wave sleep and rapid eye movement sleep to emotional item and source memory. Neurobiol. Learn. Mem. 122, 122-130. https://doi.org/10.1016/j.nlm.2014.08.013. source

In the paper

“other studies revealed comparable benefits from sleep for neutral over emotional stimuli.”

What the sources say

• Supported

  The Direct associative memory section reports no significant interaction with valence, indicating that sleep similarly benefited memory for negative and neutral information. The Discussion likewise reports sleep benefits regardless of valence, rather than a preference for emotional stimuli. (LLM, conf=0.91)

  cites [26] 26. Huguet, M., Payne, J.D., Kim, S.Y., and Alger, S.E. (2019). Overnight sleep benefits both neutral and negative direct associative and relational memory. Cogn. Affect. Behav. Neurosci. 19, 1391-1403. https://doi.org/ 10.3758/s13415-019-00746-8. source
• Supported

  In “Results from Studies Examining Sleep/Wake Contrasts” (page unavailable), the review reports that two meta-analyses found no overall evidence that sleep benefits emotional memories more than neutral ones. Its study table also lists sleep effects without sleep-by-emotion interactions, consistent with comparable benefits rather than a general emotional-memory advantage. (LLM, conf=0.87)

  cites [27] 27. Davidson, P., Jo ¨nsson, P., Carlsson, I., and Pace-Schott, E. (2021). Does sleep selectively strengthen certain memories over others based on emotion and perceived future relevance? Nat. Sci. Sleep 13, 1257- 1306. https://doi.org/10.2147/NSS.S286701. source

In the paper

“Across experimental designs, post-encoding sleep has been consistently found to enhance memory retrieval on non-verbal (e.g., object-location associations) and verbal (e.g., lists of word pairs) tasks assumed to assess hippocampus-dependent episodic types of memories.”

What the sources say

• Supported

  The section on selectivity of memory consolidation during sleep (page unavailable) reports sleep-associated recall benefits for both visual-spatial and verbal paired associates, and also describes picture-location associations. The section on reorganization of declarative memories reports improved word-pair memory at a later retrieval test. Together, these passages support the claim’s generalization across the two task types. (LLM, conf=0.90)

  cites [3] 3. Rasch, B., and Born, J. (2013). About sleep's role in memory. Physiol. Rev. 93, 681-766. https://doi.org/10.1152/physrev.00032.2012. source
• Partly supported

  The Introduction (page not supplied) says sleep following encoding appears to benefit hippocampus-dependent memory. The Results report better recognition of pictured objects after sleep than after wakefulness. The paper does not establish the claim across verbal and non-verbal tasks or experimental designs; its experiment did not test word pairs or object-location associations. (LLM, conf=0.86)

  cites [13] 13. Hanert, A., Weber, F.D., Pedersen, A., Born, J., and Bartsch, T. (2017). Sleep in humans stabilizes pattern separation performance. J. Neurosci. 37, 12238-12246. https://doi.org/10.1523/JNEUROSCI.1189-17.2017. sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites [14] 14. Zhang, J., Whitehurst, L.N., and Mednick, S.C. (2022). The role of sleep for episodic memory consolidation: stabilizing or rescuing? Neurobiol. Learn. Mem. 191, 107621. https://doi.org/10.1016/j.nlm.2022.107621. source

In the paper

“REM sleep and associated theta activity was most consistently found to be involved in the consolidation of emotional memories (e.g., fear conditioning 70,71 ) including social memory for a conspecific mouse.”

What the sources say

• Partly supported

  The Abstract and Discussion link REM-sleep theta activity to emotional memory consolidation in a fear-conditioning study. The Methods identify the subjects as rats, and the paper does not address social memory for a conspecific mouse. It also does not establish that REM sleep and theta activity are the most consistently implicated factors across emotional-memory studies. (LLM, conf=0.88)

  cites [70] 70. Totty, M.S., Chesney, L.A., Geist, P.A., and Datta, S. (2017). Sleep-depen- dent oscillatory synchronization: A role in fear memory consolidation. Front. Neural Circuits 11, 49. https://doi.org/10.3389/fncir.2017.00049. sourceExperimental — accuracy not yet measured
• Partly supported

  The Results section on fear memory expression and the Discussion link REM-sleep theta activity to contextual fear memory in rats. The paper does not study social memory for a conspecific mouse or establish that emotional memories are the memory type most consistently associated with REM sleep and theta activity. Page numbers are unavailable in our copy. (LLM, conf=0.86)

  cites [71] 71. de Almeida-Filho, D.G., Koike, B.D.V., Billwiller, F., Farias, K.S., de Sales, I.R.P., Luppi, P.H., Ribeiro, S., and Queiroz, C.M. (2021). Hippocampus- retrosplenial cortex interaction is increased during phasic REM and con- tributes to memory consolidation. Sci. Rep. 11, 13078. https://doi.org/ 10.1038/s41598-021-91659-5. sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites [72] 72. Qin, H., Fu, L., Jian, T., Jin, W., Liang, M., Li, J., Chen, Q., Yang, X., Du, H., Liao, X., et al. (2022). REM sleep-active hypothalamic neurons may contribute to hippocampal social-memory consolidation. Neuron 110, 4000-4014.e6. https://doi.org/10.1016/j.neuron.2022.09.004. source

In the paper

“the hippocampus often enters REM sleep while cortical areas still remain in SWS”

What the sources say

• ?Not checked — no full text of the sourcecites [9] 9. Dura ´n, E., Oyanedel, C.N., Niethard, N., Inostroza, M., and Born, J. (2018). Sleep stage dynamics in neocortex and hippocampus. Sleep 41, zsy060. https://doi.org/10.1093/sleep/zsy060. source
• Partly supported

  The paper reports that, in rats, the hippocampus often entered REM before the neocortex. But the illustrated simultaneous state, described in “The Hippocampus and Neocortex Can Simultaneously Exist in Different Sleep States” and Figure 3 (page unavailable), is hippocampal REM with neocortical transition-to-REM (TR), not neocortical SWS. The claim therefore overstates which cortical state remained when hippocampal REM began. (LLM, conf=0.88)

  cites [10] 10. Emrick, J.J., Gross, B.A., Riley, B.T., and Poe, G.R. (2016). Different simultaneous sleep states in the hippocampus and neocortex. Sleep 39, 2201-2209. https://doi.org/10.5665/sleep.6326. sourceExperimental — accuracy not yet measured

In the paper

“in comparison with post-encoding wakefulness, sleep increased the recruitment of neocortical networks and functional connectivity between prefrontal cortex and hippocampus at a later retrieval test, especially with declarative types of memory.”

What the sources say

• ?Not checked — no full text of the sourcecites [33] 33. Gais, S., Albouy, G., Boly, M., Dang-Vu, T.T., Darsaud, A., Desseilles, M., Rauchs, G., Schabus, M., Sterpenich, V., Vandewalle, G., et al. (2007). Sleep transforms the cerebral trace of declarative memories. Proc. Natl. Acad. Sci. USA 104, 18778-18783. https://doi.org/10.1073/pnas. 0705454104. source
• Partly supported

  The Results section ('Overnight fast spindle density and next-day functional connectivity correlations: anterior hippocampus'; page unavailable) links greater overnight spindle density to stronger hippocampal–vmPFC connectivity when participants restudied object–word pairs learned before sleep. But it reports no significant overall connectivity difference between the presleep and morning lists. The comparison was with pairs encoded after sleep, not a matched post-encoding wakefulness condition, and the neural activity was measured during restudy rather than a later retrieval test; the cited passages also do not establish increased neocortical recruitment generally. (LLM, conf=0.88)

  cites [34] 34. Cowan, E., Liu, A., Henin, S., Kothare, S., Devinsky, O., and Davachi, L. (2020). Sleep spindles promote the restructuring of memory representa- tions in ventromedial prefrontal cortex through enhanced hippocampal- cortical functional connectivity. J. Neurosci. 40, 1909-1919. https://doi. org/10.1523/JNEUROSCI.1946-19.2020. sourceExperimental — accuracy not yet measured

In the paper

“inhibiting V1 engram cells during sleep induced a more generalized fear response irrespective of the visual cue presented.”

What the sources say

• Supported

  The Results (Fig. 7) report that mice with sleep-targeted inhibition of V1 engram neurons froze more to both visual cues than controls, which the authors describe as generalization. The Results also note a qualification: mice conditioned with Y° as the shock cue still discriminated between the cues. (LLM, conf=0.91)

  cites [87] 87. Clawson, B.C., Pickup, E.J., Ensing, A., Geneseo, L., Shaver, J., Gonza- lez-Amoretti, J., Zhao, M., York, A.K., Kuhn, F.R., Swift, K., et al. (2021). Causal role for sleep-dependent reactivation of learning-activated sen- sory ensembles for fear memory consolidation. Nat. Commun. 12, 1200. https://doi.org/10.1038/s41467-021-21471-2. source

In the paper

“Multivariate pattern analyses have provided robust evidence that TMR actually induces a behaviorally relevant reactivation of cue-associated target information.”

What the sources say

• ?Not checked — no full text of the sourcecites [117] 117. Aston-Jones, G., and Bloom, F.E. (1981). Activity of norepinephrine-con- taining locus coeruleus neurons in behaving rats anticipates fluctuations in the sleep-waking cycle. J. Neurosci. 1, 876-886. https://doi.org/10. 1523/JNEUROSCI.01-08-00876.1981. 207. Berridge, C.W. (2008). Noradrenergic modulation of arousal. Brain Res. Rev. 58, 1-17. https://doi.org/10.1016/j.brainresrev.2007.10.013. 208. Rodenkirch, C., Liu, Y., Schriver, B.J., and Wang, Q. (2019). Locus coeru- leus activation enhances thalamic feature selectivity via norepinephrine regulation of intrathalamic circuit dynamics. Na source
• ?Not checked — no full text of the sourcecites [119] 119. Hobson, J.A. (2010). Sleeping, waking, and dreaming. In Encyclopedia of Behavioral Neuroscience (Elsevier Inc.), pp. 237-256. 221. source
• ?Could not verify

  The Results passages concern sleep-spindle measurements and associations, while Methods > Topographical spindle analyses describes identifying multivariate outliers, not multivariate pattern analysis of memory reactivation. Searches for TMR, cue-associated information, and behaviorally relevant reactivation found no passage in our copy addressing the claim. (LLM, conf=0.77)

  cites [127] 127. Purcell, S.M., Manoach, D.S., Demanuele, C., Cade, B.E., Mariani, S., Cox, R., Panagiotaropoulou, G., Saxena, R., Pan, J.Q., Smoller, J.W., et al. (2017). Characterizing sleep spindles in 11,630 individuals from the National Sleep Research Resource. Nat. Commun. 8, 15930. https://doi.org/10.1038/ncomms15930. 253. sourceExperimental — accuracy not yet measured

In the paper

“several studies identified a causal relationship between reactivation and memory consolidation.”

What the sources say

• Partly supported

  The Discussion (page not provided) reports that reactivation predicts later spatial-memory performance and describes a functional link. It does not establish the claimed causal relationship: the authors say experimental evidence has been largely lacking and note that studies disrupting sharp-wave ripples could not rule out effects on consolidation through mechanisms other than reactivation. (LLM, conf=0.88)

  cites [83] 83. Dupret, D., O'Neill, J., Pleydell-Bouverie, B., and Csicsvari, J. (2010). The reorganization and reactivation of hippocampal maps predict spatial memory performance. Nat. Neurosci. 13, 995-1002. https://doi.org/10. 1038/nn.2599. sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites [84] 84. Girardeau, G., Benchenane, K., Wiener, S.I., Buzsa ´ki, G., and Zugaro, M.B. (2009). Selective suppression of hippocampal ripples impairs spatial mem- ory. Nat. Neurosci. 12, 1222-1223. https://doi.org/10.1038/nn.2384. source
• Partly supported

  The Discussion reports that this study directly disrupted SWR reactivation and found impaired later reinstatement of newly formed hippocampal assembly patterns; it interprets this, together with earlier behavioral findings, as evidence that reactivation supports memory consolidation. But the Introduction and Discussion say earlier studies had not established whether their effects were caused by disrupted reactivation. This source therefore supports a causal role for reactivation in stabilizing the patterns, not the claim that several studies had identified that causal relationship. (LLM, conf=0.85)

  cites [85] 85. van de Ven, G.M., Trouche, S., McNamara, C.G., Allen, K., and Dupret, D. (2016). Hippocampal offline reactivation consolidates recently formed cell assembly patterns during sharp wave-ripples. Neuron 92, 968-974. https://doi.org/10.1016/j.neuron.2016.10.020. sourceExperimental — accuracy not yet measured

In the paper

“intracranial recordings have provided corroborative evidence for stimulus-specific and sequential reactivations during human sleep.”

What the sources say

• ?Not checked — no full text of the sourcecites [110] 110. Wierzynski, C.M., Lubenov, E.V., Gu, M., and Siapas, A.G. (2009). State- dependent spike-timing relationships between hippocampal and pre- frontal circuits during sleep. Neuron 61, 587-596. https://doi.org/10. 1016/j.neuron.2009.01.011. 170. Eschenko, O., Magri, C., Panzeri, S., and Sara, S.J. (2012). Noradren- ergic neurons of the locus coeruleus are phase locked to cortical up- down states during sleep. Cereb. Cortex 22, 426-435. https://doi.org/ 10.1093/cercor/bhr121. 171. source
• Partly supported

  The Results report intracranial recordings during human sleep, and the Discussion interprets the findings as supporting hippocampal information reactivation and transfer. The paper does not report stimulus-specific or sequential reactivation of learned content; the Discussion notes that the study obtained no concomitant behavior. (LLM, conf=0.85)

  cites [112] 112. Helfrich, R.F., Lendner, J.D., Mander, B.A., Guillen, H., Paff, M., Mnatsa- kanyan, L., Vadera, S., Walker, M.P., Lin, J.J., and Knight, R.T. (2019). Bidirectional prefrontal-hippocampal dynamics organize information transfer during sleep in humans. Nat. Commun. 10, 3572. https://doi. org/10.1038/s41467-019-11444-x. 187. sourceExperimental — accuracy not yet measured

In the paper

“offline memory reactivations and recently also signs of sequential replay 104,105 have been observed with noninvasive neuroimaging in humans.”

What the sources say

• Partly supported

  The Introduction (page unavailable) says human neuroimaging studies show reactivation of learning-related brain regions during post-encoding sleep. It describes reinstatement of neuronal firing sequences in rodents, not noninvasive observation of sequential replay in humans. Thus the reactivation portion is supported, but the human sequential-replay portion is not. (LLM, conf=0.86)

  cites [104] 104. Cousins, J.N., El-Deredy, W., Parkes, L.M., Hennies, N., and Lewis, P.A. (2014). Cued memory reactivation during slow-wave sleep promotes explicit knowledge of a motor sequence. J. Neurosci. 34, 15870- 15876. https://doi.org/10.1523/JNEUROSCI.1011-14.2014. 125. Nicolas, J., King, B.R., Levesque, D., Lazzouni, L., Coffey, E., Swinnen, S., Doyon, J., Carrier, J., and Albouy, G. (2022). Sigma oscillations pro- tect or reinstate motor memory depending on their temporal coordination with slow waves. eLife 11, e73930. https://doi.org/10.7554/eLife.73930. 126. van Dongen, E.V., Takashima, A., Ba sourceExperimental — accuracy not yet measured
• Partly supported

  The Abstract reports offline reactivation of activity patterns in rat hippocampal recordings, and the Discussion mentions evidence of sequence reactivation during slow-wave sleep. Neither passage supports the claim that these observations were made in humans using noninvasive neuroimaging; the paper describes implanted tetrode recordings in rats (Recording protocol). (LLM, conf=0.88)

  cites [105] 105. Batterink, L.J., Creery, J.D., and Paller, K.A. (2016). Phase of sponta- neous slow oscillations during sleep influences memory-related process- ing of auditory cues. J. Neurosci. 36, 1401-1409. https://doi.org/10. 1523/JNEUROSCI.3175-15.2016. 130. Bendor, D., and Wilson, M.A. (2012). Biasing the content of hippocampal replay during sleep. Nat. Neurosci. 15, 1439-1444. https://doi.org/10. 1038/nn.3203. 131. Kudrimoti, H.S., Barnes, C.A., and McNaughton, B.L. (1999). Reactiva- tion of hippocampal cell assemblies: effects of behavioral state, experi- ence, and EEG dynamics. J. Neurosci. 19, sourceExperimental — accuracy not yet measured

In the paper

“Using machine learning classifiers, categories of previously encoded stimuli (e.g., faces vs. houses; objects vs. scenes) could be decoded solely from sleep EEG, revealing that category-specific information is reactivated during sleep.”

What the sources say

• Partly supported

  The ‘Sleep and Structural Synaptic Changes’ section reports reactivation of motor-task-related neurons in mice during sleep, supporting the general idea of sleep reactivation. Neither that section nor the EEG discussion in ‘Sleep’s Effect on Neuronal Excitability’ reports machine-learning decoding of previously encoded stimulus categories from sleep EEG; searches for those methods and categories found no such passage. Page numbers are unavailable in our copy. (LLM, conf=0.82)

  cites [113] 113. Niethard, N., Burgalossi, A., and Born, J. (2017). Plasticity during sleep is linked to specific regulation of cortical circuit activity. Front. Neural Cir- cuits 11, 65. https://doi.org/10.3389/fncir.2017.00065. 188. Schreiner, T., Kaufmann, E., Noachtar, S., Mehrkens, J.H., and Staudigl, T. (2022). The human thalamus orchestrates neocortical oscillations dur- ing NREM sleep. Nat. Commun. 13, 5231. https://doi.org/10.1038/ s41467-022-32840-w. 189. Alizadeh, Z., Azimi, A., and Ghorbani, M. (2022). Enhancement of hippo- campal-thalamocortical temporal coordination during slow-frequency lon sourceExperimental — accuracy not yet measured
• ?Could not verify

  Our copy reports NeuroGrid and hippocampal recordings in rats and stronger ripple coupling during sleep after spatial learning (Abstract; page unavailable). Searches for machine-learning classification, stimulus categories, and decoding from sleep EEG found no passage addressing those claims. (LLM, conf=0.76)

  cites [114] 114. Khodagholy, D., Gelinas, J.N., and Buzsa ´ki, G. (2017). Learning- enhanced coupling between ripple oscillations in association cortices and hippocampus. Science 358, 369-372. 192. sourceExperimental — accuracy not yet measured

In the paper

“There is, however, also evi-dence for hippocampus-independent replay in extrahippocampal sites, e.g., in superficial layers of medial entorhinal cortex and in prefrontal cortex.”

What the sources say

• ?Not checked — no full text of the sourcecites [91] 91. O'Neill, J., Boccara, C.N., Stella, F., Schoenenberger, P., and Csicsvari, J. (2017). Superficial layers of the medial entorhinal cortex replay inde- pendently of the hippocampus. Science 355, 184-188. https://doi.org/ 10.1126/science.aag2787. source
• Partly supported

  The Discussion reports that superficial-layer medial entorhinal grid cells rarely reactivate in coordination with hippocampal place cells, and the References identify a study of replay independent of the hippocampus in those layers. For medial prefrontal cortex, however, the Discussion describes activity correlated with hippocampal activity during replay, not hippocampus-independent replay. No page numbers are provided in our copy. (LLM, conf=0.85)

  cites [93] 93. Trettel, S.G., Trimper, J.B., Hwaun, E., Fiete, I.R., and Colgin, L.L. (2019). Grid cell co-activity patterns during sleep reflect spatial overlap of grid fields during active behaviors. Nat. Neurosci. 22, 609-617. https://doi. org/10.1038/s41593-019-0359-6. sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites [98] 98. Kaefer, K., Nardin, M., Blahna, K., and Csicsvari, J. (2020). Replay of behavioral sequences in the medial prefrontal cortex during rule switch- ing. Neuron 106, 154-165.e6. https://doi.org/10.1016/j.neuron.2020. 01.015. source

In the paper

“The beneficial effect of TMR during non-REM sleep on memory performance is well established across a variety of memory domains, 115 including visuospatial 116-119 and verbal 120,121 declarative memory as well as motor learning.”

What the sources say

• ?Could not verify

  The Discussion describes spontaneous ripple coupling during NREM sleep and a possible role in memory consolidation. The Results concern cued recall while patients were awake, not targeted memory reactivation during sleep. Searches for TMR and for the specified memory domains found no passage establishing a benefit across those domains. (LLM, conf=0.76)

  cites [115] 115. Dickey, C.W., Verzhbinsky, I.A., Jiang, X., Rosen, B.Q., Kajfez, S., Stede- lin, B., Shih, J.J., Ben-Haim, S., Raslan, A.M., Eskandar, E.N., et al. (2022). Widespread ripples synchronize human cortical activity during sleep, waking, and memory recall. Proc. Natl. Acad. Sci. USA 119. e2107797119. https://doi.org/10.1073/pnas.2107797119. 193. Ferna ´ndez-Ruiz, A., Oliva, A., Fermino de Oliveira, E.F., Rocha-Almeida, F., Tingley, D., and Buzsa ´ki, G. (2019). Long-duration hippocampal sharp wave ripples improve memory. Science 364, 1082-1086. https://doi.org/ 10.1126/science.aax0758. 194. Ol sourceExperimental — accuracy not yet measured
• ?Could not verify

  The Introduction says NREM sleep facilitates memory consolidation, and the Discussion reports that externally facilitating slow oscillations can boost later retrieval. Neither passage addresses targeted memory reactivation (TMR) or establishes its benefits across memory domains. Searches for TMR, reactivation cues, and the named domains found no passage that verifies the attributed claim. (LLM, conf=0.72)

  cites [116] 116. Staresina, B.P., Bergmann, T.O., Bonnefond, M., van der Meij, R., Jen- sen, O., Deuker, L., Elger, C.E., Axmacher, N., and Fell, J. (2015). Hierar- chical nesting of slow oscillations, spindles and ripples in the human hip- pocampus during sleep. Nat. Neurosci. 18, 1679-1686. https://doi.org/ 10.1038/nn.4119. 198. Mo ¨lle, M., and Born, J. (2011). Slow oscillations orchestrating fast oscil- lations and memory consolidation. In Prog. Brain Res., 193, pp. 93-110. 199. Oyanedel, C.N., Dura ´n, E., Niethard, N., Inostroza, M., and Born, J. (2020). Temporal associations between sleep slow osci sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites [117] 117. Aston-Jones, G., and Bloom, F.E. (1981). Activity of norepinephrine-con- taining locus coeruleus neurons in behaving rats anticipates fluctuations in the sleep-waking cycle. J. Neurosci. 1, 876-886. https://doi.org/10. 1523/JNEUROSCI.01-08-00876.1981. 207. Berridge, C.W. (2008). Noradrenergic modulation of arousal. Brain Res. Rev. 58, 1-17. https://doi.org/10.1016/j.brainresrev.2007.10.013. 208. Rodenkirch, C., Liu, Y., Schriver, B.J., and Wang, Q. (2019). Locus coeru- leus activation enhances thalamic feature selectivity via norepinephrine regulation of intrathalamic circuit dynamics. Na source
• ?Not checked — no full text of the sourcecites [118] 118. Sara, S.J. (2015). Locus Coeruleus in time with the making of memories. Curr. Opin. Neurobiol. 35, 87-94. https://doi.org/10.1016/j.conb.2015. 07.004. 210. Tronel, S., Feenstra, M.G., and Sara, S.J. (2004). Noradrenergic action in prefrontal cortex in the late stage of memory consolidation. Learn. Mem. 11, 453-458. https://doi.org/10.1101/lm.74504. 211. Menon, J.M.L., Nolten, C., Achterberg, E.J.M., Joosten, R.N.J.M.A., De- matteis, M., Feenstra, M.G.P., Drinkenburg, W.H.P., and Leenaars, C.H.C. (2019). Brain microdialysate monoamines in relation to circadian rhythms, sleep, and sleep dep source
• ?Not checked — no full text of the sourcecites [119] 119. Hobson, J.A. (2010). Sleeping, waking, and dreaming. In Encyclopedia of Behavioral Neuroscience (Elsevier Inc.), pp. 237-256. 221. source
• ?Not checked — no full text of the sourcecites [120] 120. Boyce, R., Glasgow, S.D., Williams, S., and Adamantidis, A. (2016). Causal evidence for the role of REM sleep theta rhythm in contextual memory consolidation. Science 352, 812-816. https://doi.org/10.1126/ science.aad5252. 222. source
• Partly supported

  In “Synaptic renormalization by down-selection and memory” (page unavailable), the review reports experimental benefits of cuing during sleep and discusses selective enhancement in a model of declarative learning. It does not establish that targeted memory reactivation during non-REM sleep benefits performance across the range of memory domains claimed; its discussion of visuomotor learning concerns sleep benefits generally, not cuing. (LLM, conf=0.84)

  cites [121] 121. Izawa, S., Chowdhury, S., Miyazaki, T., Mukai, Y., Ono, D., Inoue, R., Oh- mura, Y., Mizoguchi, H., Kimura, K., Yoshioka, M., et al. (2019). REM sleep-active MCH neurons are involved in forgetting hippocampus- dependent memories. Science 365, 1308-1313. https://doi.org/10. 1126/science.aax9238. 223. Tononi, G., and Cirelli, C. (2014). Sleep and the price of plasticity: from synaptic and cellular homeostasis to memory consolidation and integra- tion. Neuron 81, 12-34. https://doi.org/10.1016/j.neuron.2013.12.025. 224. Tononi, G., and Cirelli, C. (2003). Sleep and synaptic homeostasis: a hy sourceExperimental — accuracy not yet measured

In the paper

“Inducing reactivations of the V1 engram cells during sleep after conditioning enhanced the formation of a cue-specific fear memory.”

What the sources say

• Partly supported

  The Introduction and Results (Fig. 7; page unavailable) report that V1 engram neurons reactivate during post-conditioning sleep and that inhibiting them during sleep disrupts cue-specific fear memory consolidation. The paper does not report that experimentally inducing reactivation during sleep enhanced fear memory; its optogenetic activation experiments assessed other outcomes or conditions. (LLM, conf=0.88)

  cites [87] 87. Clawson, B.C., Pickup, E.J., Ensing, A., Geneseo, L., Shaver, J., Gonza- lez-Amoretti, J., Zhao, M., York, A.K., Kuhn, F.R., Swift, K., et al. (2021). Causal role for sleep-dependent reactivation of learning-activated sen- sory ensembles for fear memory consolidation. Nat. Commun. 12, 1200. https://doi.org/10.1038/s41467-021-21471-2. sourceExperimental — accuracy not yet measured

In the paper

“memory reactivation is a far more ubiquitous phenomenon, as it is observed in many brain regions, ranging from early sensory [87][88][89] and motor cortices, 65,90 over medial entorhinal cortex [91][92][93] and higher-order association areas like the parietal [94][95][96] and prefrontal cortices [97][98][99] to different subcortical areas.”

What the sources say

• Supported

  The citation is attached to the sentence’s early-sensory-cortex example, not to its full list of brain regions. The Results section reports reactivation of orientation-selective primary visual cortex (V1) ensembles during sleep after visually cued fear conditioning in mice. The Introduction also notes that reactivation has been observed in multiple brain regions. (LLM, conf=0.93)

  cites [87] 87. Clawson, B.C., Pickup, E.J., Ensing, A., Geneseo, L., Shaver, J., Gonza- lez-Amoretti, J., Zhao, M., York, A.K., Kuhn, F.R., Swift, K., et al. (2021). Causal role for sleep-dependent reactivation of learning-activated sen- sory ensembles for fear memory consolidation. Nat. Commun. 12, 1200. https://doi.org/10.1038/s41467-021-21471-2. source
• ?Not checked — no full text of the sourcecites [88] 88. Ji, D., and Wilson, M.A. (2007). Coordinated memory replay in the visual cortex and hippocampus during sleep. Nat. Neurosci. 10, 100-107. https://doi.org/10.1038/nn1825. source
• Supported

  The Results section reports significant reactivation of waking experience in auditory cortex during sleep. This supports the sensory-cortex portion of the citing sentence, where this paper is cited; the other regions are accompanied by separate citations. (LLM, conf=0.90)

  cites [89] 89. Rothschild, G., Eban, E., and Frank, L.M. (2017). A cortical-hippocampal- cortical loop of information processing during memory consolidation. Nat. Neurosci. 20, 251-259. https://doi.org/10.1038/nn.4457. source

In the paper

“parallel recordings from both hippocampus and cortical or subcortical regions revealed a close temporal coordination of reactivation across sites 92,100,102”

What the sources say

• Supported

  The paper reports concurrent recordings from hippocampal CA1 and the cortical medial entorhinal cortex (p. 1). It finds that grid-cell activity during rest replay peaked about 10 ms after place-cell activity (p. 1) and concludes that the two were closely coordinated during replay (p. 3). (LLM, conf=0.92)

  cites [92] 92. O ´lafsdo ´ttir, H.F., Carpenter, F., and Barry, C. (2016). Coordinated grid and place cell replay during rest. Nat. Neurosci. 19, 792-794. https:// doi.org/10.1038/nn.4291. source
• ?Not checked — no full text of the sourcecites [100] 100. Maquet, P., Laureys, S., Peigneux, P., Fuchs, S., Petiau, C., Phillips, C., Aerts, J., Del Fiore, G., Degueldre, C., Meulemans, T., et al. (2000). Expe- rience-dependent changes in cerebral activation during human REM sleep. Nat. Neurosci. 3, 831-836. https://doi.org/10.1038/77744. 107. Peigneux, P., Laureys, S., Fuchs, S., Collette, F., Perrin, F., Reggers, J., Phil- lips, C., Degueldre, C., Del Fiore, G., Aerts, J., et al. (2004). Are spatial mem- ories strengthened in the human hippocampus during slow wave sleep? Neuron 44, 535-545. https://doi.org/10.1016/j.neuron.2004.10.007. 108. Be source
• ?Could not verify

  The Online Methods, ‘Sleep Recording’ section (page not provided) describes scalp EEG recordings, not parallel recordings from the hippocampus and cortical or subcortical sites. The copy does not report temporal coordination of reactivation across those sites; searches for parallel recordings and hippocampal–cortical coordination found no such passage. (LLM, conf=0.75)

  cites [102] 102. Antony, J.W., Gobel, E.W., O'Hare, J.K., Reber, P.J., and Paller, K.A. (2012). Cued memory reactivation during sleep influences skill learning. Nat. Neurosci. 15, 1114-1116. https://doi.org/10.1038/nn.3152. 117. Cairney, S.A., Guttesen, A.A ´.V., El Marj, N., and Staresina, B.P. (2018). Memory consolidation is linked to spindle-mediated information pro- cessing during sleep. Curr. Biol. 28, 948-954.e4. https://doi.org/10. 1016/j.cub.2018.01.087. 118. Rasch, B., B€ uchel, C., Gais, S., and Born, J. (2007). Odor cues during slow-wave sleep prompt declarative memory consolidation. Science 31 sourceExperimental — accuracy not yet measured

In the paper

“In both of these studies, the accuracy of the classifier to correctly identify the previously encoded stimulus category was positively correlated to later memory recall”

What the sources say

• ?Could not verify

  The Conclusion discusses sleep-related changes in cortical activity and their possible role in strengthening memories, but does not report classifier results. Searches for classifier accuracy, encoded stimulus categories, and correlations with later recall found no passage in our copy addressing the claimed finding in either study. (LLM, conf=0.74)

  cites [113] 113. Niethard, N., Burgalossi, A., and Born, J. (2017). Plasticity during sleep is linked to specific regulation of cortical circuit activity. Front. Neural Cir- cuits 11, 65. https://doi.org/10.3389/fncir.2017.00065. 188. Schreiner, T., Kaufmann, E., Noachtar, S., Mehrkens, J.H., and Staudigl, T. (2022). The human thalamus orchestrates neocortical oscillations dur- ing NREM sleep. Nat. Commun. 13, 5231. https://doi.org/10.1038/ s41467-022-32840-w. 189. Alizadeh, Z., Azimi, A., and Ghorbani, M. (2022). Enhancement of hippo- campal-thalamocortical temporal coordination during slow-frequency lon sourceExperimental — accuracy not yet measured
• ?Could not verify

  The study description and results (indexed under Abstract; page unavailable) concern rats learning reward locations and increased hippocampal–cortical ripple coupling after training. Searches for classifier accuracy, encoded stimulus categories, and subsequent recall found no passage reporting the claimed correlation. (LLM, conf=0.75)

  cites [114] 114. Khodagholy, D., Gelinas, J.N., and Buzsa ´ki, G. (2017). Learning- enhanced coupling between ripple oscillations in association cortices and hippocampus. Science 358, 369-372. 192. sourceExperimental — accuracy not yet measured

In the paper

“The reactivation of task-relevant regions was found to be particular prominent during sleep oscillatory events linked to memory consolidation, such as spindles and SOs.”

What the sources say

• Partly supported

  The Discussion (page not provided) reports coordinated memory reactivation in the hippocampus and visual cortex during sleep up-states associated with slow oscillations. It does not say that reactivation was particularly prominent during spindles, or establish the claimed prominence across both types of event. (LLM, conf=0.86)

  cites [108] 108. Davidson, T.J., Kloosterman, F., and Wilson, M.A. (2009). Hippocampal replay of extended experience. Neuron 63, 497-507. https://doi.org/10. 1016/j.neuron.2009.07.027. 149. O ´lafsdo ´ttir, H.F., Barry, C., Saleem, A.B., Hassabis, D., and Spiers, H.J. (2015). Hippocampal place cells construct reward related sequences through unexplored space. eLife 4, e06063. https://doi.org/10.7554/eL- ife.06063. 150. Stella, F., Baracskay, P., O'Neill, J., and Csicsvari, J. (2019). Hippocam- pal reactivation of random trajectories resembling brownian diffusion. Neuron 102, 450-461.e7. e457. https://doi. sourceExperimental — accuracy not yet measured
• Partly supported

  The Role of Sleep SWR Replay reports cortical reactivation during slow-wave sleep, and The Fidelity of Replay says prefrontal reactivation strength peaked near a transition in cortical slow oscillations. The paper mentions sleep spindles in its references, but our copy does not say that task-related reactivation was particularly prominent during spindles. (LLM, conf=0.86)

  cites [109] 109. Roumis, D.K., and Frank, L.M. (2015). Hippocampal sharp-wave ripples in waking and sleeping states. Curr. Opin. Neurobiol. 35, 6-12. https:// doi.org/10.1016/j.conb.2015.05.001. 156. Tang, W., Shin, J.D., Frank, L.M., and Jadhav, S.P. (2017). Hippocampal- prefrontal reactivation during learning is stronger in awake compared with sleep states. J. Neurosci. 37, 11789-11805. https://doi.org/10. 1523/JNEUROSCI.2291-17.2017. 157. de Sousa, A.F., Cowansage, K.K., Zutshi, I., Cardozo, L.M., Yoo, E.J., Leutgeb, S., and Mayford, M. (2019). Optogenetic reactivation of memory ensembles in the retros sourceExperimental — accuracy not yet measured

In the paper

“In general, reactivation and replay in both wake and sleep states display largely similar features.”

What the sources say

• Supported

  Section 8 (Conclusions; page not supplied) says that little evidence separates the phenomenology of awake and sleep replay and that activity patterns appear strongly conserved. It acknowledges quantitative differences, which is consistent with the claim's qualifier “largely.” (LLM, conf=0.94)

  cites [140] source

In the paper

“Signs of reward-related biasing of reactivations during sleep and wake have been likewise observed in humans using fMRI.”

What the sources say

• ?Not checked — no full text of the sourcecites [111] 111. Adamantidis, A.R., Gutierrez Herrera, C., and Gent, T.C. (2019). Oscil- lating circuitries in the sleeping brain. Nat. Rev. Neurosci. 20, 746-762. https://doi.org/10.1038/s41583-019-0223-4. 172. Fellin, T., Halassa, M.M., Terunuma, M., Succol, F., Takano, H., Frank, M., Moss, S.J., and Haydon, P.G. (2009). Endogenous nonneuronal mod- ulators of synaptic transmission control cortical slow oscillations in vivo. Proc. Natl. Acad. Sci. USA 106, 15037-15042. https://doi.org/10.1073/ pnas.0906419106. 173. Crunelli, V., and Hughes, S.W. (2010). The slow (<1 Hz) rhythm of non- REM sleep: a dialog source
• Partly supported

  The Results report preferential reactivation of high-reward contexts in human hippocampal fMRI data during post-learning rest, supporting the wakeful-rest portion of the claim. The Discussion’s references to sleep concern rodent replay and human intracranial recordings, not reward-biased reactivation during sleep measured with human fMRI. Page numbers are unavailable in our copy. (LLM, conf=0.90)

  cites [147] sourceExperimental — accuracy not yet measured

In the paper

“TMR has also been used in rodent studies, confirming that auditory cues, which had been paired to specific trajectories dur-ing a spatial learning task, enhance hippocampal reactivation of the associated trajectory when delivered during subsequent sleep.”

What the sources say

• Partly supported

  The Results describe rats learning a sound-guided Y-track task and show that sounds delivered during sleep biased auditory-cortex patterns that predicted subsequent hippocampal CA1 activity. They do not establish enhanced reactivation of the trajectory associated with a cue: the Results state that sound identity could not be significantly decoded from CA1 ensembles. The Discussion distinguishes this paper’s finding from a previous study showing that sleep sounds can bias hippocampal reactivation. (LLM, conf=0.88)

  cites [89] 89. Rothschild, G., Eban, E., and Frank, L.M. (2017). A cortical-hippocampal- cortical loop of information processing during memory consolidation. Nat. Neurosci. 20, 251-259. https://doi.org/10.1038/nn.4457. sourceExperimental — accuracy not yet measured
• ?Could not verify

  The Introduction and Discussion discuss memory during infant sleep, but do not describe rodent targeted memory reactivation. Searches for auditory cues paired with spatial trajectories and for rodent targeted memory reactivation found no passage addressing the claimed hippocampal reactivation effect. The retrieved passages have no page numbers. (LLM, conf=0.75)

  cites [130] 130. Friedrich, M., Mo ¨lle, M., Friederici, A.D., and Born, J. (2020). Sleep- dependent memory consolidation in infants protects new episodic mem- ories from existing semantic memories. Nat. Commun. 11, 1298. https:// doi.org/10.1038/s41467-020-14850-8. 271. sourceExperimental — accuracy not yet measured

In the paper

“awake memory reactivations have been shown to enhance subsequent recall performance.”

What the sources say

• Partly supported

  The Results section (‘Reactivation of waking firing patterns during rest’; page unavailable) reports that goal-related reactivation during rest predicted subsequent spatial-memory performance, and the Discussion says reactivation predicted how well a location was remembered. The paper does not show that awake reactivations themselves enhanced recall: the reported relationship is predictive, and the rest-period events are not identified as exclusively awake. (LLM, conf=0.86)

  cites [83] 83. Dupret, D., O'Neill, J., Pleydell-Bouverie, B., and Csicsvari, J. (2010). The reorganization and reactivation of hippocampal maps predict spatial memory performance. Nat. Neurosci. 13, 995-1002. https://doi.org/10. 1038/nn.2599. sourceExperimental — accuracy not yet measured
• Partly supported

  In “Linking awake replay and memory consolidation” (page unavailable), the review reports that the strength of awake reactivation predicted subsequent memory-task performance in animals. That supports an association, but not the claim that reactivation was shown to enhance subsequent recall performance: the passage does not establish a causal improvement or identify a recall measure. (LLM, conf=0.86)

  cites [151] sourceExperimental — accuracy not yet measured
• Supported

  The Discussion reports that experimentally reactivating individual study events during an active delay period was shown to bolster recall for those events. The paper's own finding is correlational, but its report of the experimental finding supports the claim. (LLM, conf=0.88)

  cites [152] source

In the paper

“This time course fits with findings in humans of preferential time windows for memory reactivations 3 and 6 h into sleep after encountering a novel visuospatial memory paradigm.”

What the sources say

• ?Could not verify

  The section on excitation/inhibition balance discusses replay of newly encoded memories during sleep, but it does not report preferential reactivation windows in humans. Searches for visuospatial learning and reactivation at 3 and 6 hours found no passage addressing those details; the copy’s relevant studies instead concern cortical activity in mice. (LLM, conf=0.74)

  cites [113] 113. Niethard, N., Burgalossi, A., and Born, J. (2017). Plasticity during sleep is linked to specific regulation of cortical circuit activity. Front. Neural Cir- cuits 11, 65. https://doi.org/10.3389/fncir.2017.00065. 188. Schreiner, T., Kaufmann, E., Noachtar, S., Mehrkens, J.H., and Staudigl, T. (2022). The human thalamus orchestrates neocortical oscillations dur- ing NREM sleep. Nat. Commun. 13, 5231. https://doi.org/10.1038/ s41467-022-32840-w. 189. Alizadeh, Z., Azimi, A., and Ghorbani, M. (2022). Enhancement of hippo- campal-thalamocortical temporal coordination during slow-frequency lon sourceExperimental — accuracy not yet measured

In the paper

“Memory reactivations occur preferentially in non-REM sleep.”

What the sources say

• ?Could not verify

  The Introduction (page unavailable) describes sleep spindles during non-REM sleep as involved in memory consolidation, and the Results relate spindle activity to later memory. Neither passage addresses memory reactivation or whether it occurs preferentially in non-REM sleep. Searches for “reactivation” and a REM-versus-non-REM comparison found no passage that settles the claim. (LLM, conf=0.73)

  cites [131] 131. Friedrich, M., Mo ¨lle, M., Born, J., and Friederici, A.D. (2022). Memory for nonadjacent dependencies in the first year of life and its relation to sleep. Nat. Commun. 13, 7896. https://doi.org/10.1038/s41467-022-35558-x. 272. Mooney, L.N., Johnson, E.G., Prabhakar, J., and Ghetti, S. (2021). Mem- ory-related hippocampal activation during sleep and temporal memory in toddlers. Dev. Cogn. Neurosci. 47, 100908. https://doi.org/10.1016/j. dcn.2020.100908. 273. Johnson, E.G., Mooney, L., Graf Estes, K., Nordahl, C.W., and Ghetti, S. (2021). Activation for newly learned words in left medial-t sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites [100] 100. Maquet, P., Laureys, S., Peigneux, P., Fuchs, S., Petiau, C., Phillips, C., Aerts, J., Del Fiore, G., Degueldre, C., Meulemans, T., et al. (2000). Expe- rience-dependent changes in cerebral activation during human REM sleep. Nat. Neurosci. 3, 831-836. https://doi.org/10.1038/77744. 107. Peigneux, P., Laureys, S., Fuchs, S., Collette, F., Perrin, F., Reggers, J., Phil- lips, C., Degueldre, C., Del Fiore, G., Aerts, J., et al. (2004). Are spatial mem- ories strengthened in the human hippocampus during slow wave sleep? Neuron 44, 535-545. https://doi.org/10.1016/j.neuron.2004.10.007. 108. Be source

In the paper

“Moreover, generalization of a specific contextual fear memory to a similar context, but not consolidation of individual contexts, critically depended on the co-reactivation of engram ensembles for both context experiences during sleep 97 (Figure 1).”

What the sources say

• Supported

  The Results section on postlearning sleep reports that synchronized reactivation of relevant stored engrams in the anterior cingulate cortex was necessary for memory assimilation but dispensable for consolidation of the new memory. The Results on contextual similarity show fear transfer from a conditioned square context to a previously experienced, geometrically similar context, while freezing in the conditioned context remained high. (LLM, conf=0.90)

  cites [97] 97. Aly, M.H., Abdou, K., Okubo-Suzuki, R., Nomoto, M., and Inokuchi, K. (2022). Selective engram coreactivation in idling brain inspires implicit learning. Proc. Natl. Acad. Sci. USA 119. e2201578119. https://doi.org/ 10.1073/pnas.2201578119. source

In the paper

“As to the classical fast spindle, accumulating evidence indicates the importance of precise phase-amplitude coupling between these spindles and SO upstates for effective consolidation.61,186”

What the sources say

• Supported

  The Results section (‘SO-spindle coupling predicts overnight memory consolidation’; page unavailable) reports that retention was greatest when spindle amplitude peaked close to the SO up-state. The Methods section identifies the analyzed spindle-amplitude range as 12–16 Hz, consistent with the claim’s fast spindles. (LLM, conf=0.91)

  cites [61] 61. Helfrich, R.F., Mander, B.A., Jagust, W.J., Knight, R.T., and Walker, M.P. (2018). Old brains come uncoupled in sleep: slow wave-spindle syn- chrony, brain atrophy, and forgetting. Neuron 97, 221-230.e4. https:// doi.org/10.1016/j.neuron.2017.11.020. source
• Supported

  The Introduction identifies the studied spindles as 12–16 Hz and says cortical slow oscillations precisely coordinate them. The Results report that spindle amplitudes are predicted by slow-oscillation phase and that spindles preferentially peak at the upstate. The Discussion reports evidence that spindle timing relative to the slow oscillation, rather than spindle activity alone, determines the success of information reactivation and consolidation (page numbers unavailable). (LLM, conf=0.90)

  cites [186] source

In the paper

“By contrast, suppression during delta wave upstates improved memory.”

What the sources say

• Supported

  The Results section (‘Effects of targeting δ wave related activity’) reports that optogenetic inhibition during δ-wave up-states produced greater task-performance gains than inhibition at control times in rats. The Discussion likewise says inhibition during δ waves increased reactivation strength and offline gains. (LLM, conf=0.94)

  cites [178] source

In the paper

“enhanced memory retention after applying noninvasive brain stimulation during sleep to induce slow waves.[50][51][52]63,175”

What the sources say

• ?Not checked — no full text of the sourcecites [50] 50. Marshall, L., Helgado ´ttir, H., Mo ¨lle, M., and Born, J. (2006). Boosting slow oscillations during sleep potentiates memory. Nature 444, 610-613. https://doi.org/10.1038/nature05278. source
• ?Not checked — no full text of the sourcecites [51] 51. Ngo, H.V., Martinetz, T., Born, J., and Mo ¨lle, M. (2013). Auditory closed- loop stimulation of the sleep slow oscillation enhances memory. Neuron 78, 545-553. https://doi.org/10.1016/j.neuron.2013.03.006. source
• Supported

  The Conclusion (page unavailable) says auditory stimulation during sleep enhances slow oscillations and that multiple experiments found a significant benefit for memory retention. It also notes that the memory effect varies by learning material and memory system; the claim is supported as a report of observed benefits, not a universal effect. (LLM, conf=0.90)

  cites [52] 52. Harrington, M.O., and Cairney, S.A. (2021). Sounding it out: auditory stimulation and overnight memory processing. Curr. Sleep Med. Rep. 7, 112-119. https://doi.org/10.1007/s40675-021-00207-0. source

In the paper

“Suppressing motor cortex activity during the SO upstate, as expected, impaired memory for the trained motor neuron response.”

What the sources say

• Supported

  The Results section reports that optogenetic inhibition during slow-oscillation up-states in post-training sleep led to significantly worse performance on the BMI task after awakening. The Methods section explains that rats learned to control the task apparatus through the activity of selected motor-cortex neurons. Page numbers were not available in our copy. (LLM, conf=0.91)

  cites [178] source

In the paper

“Generally, sleep replay appears to feature a less faithful recapitulation of previous experience compared with wake replay.137,[154][155][156]”

What the sources say

• Supported

  The Discussion states that replay in waking and more sleep-like states was of higher fidelity in the awake state when the animal had recently been in motion. It also describes sleep and sleep-like activity as a less faithful recapitulation of experience than waking events, supporting the citing sentence’s hedged comparison. (LLM, conf=0.93)

  cites [137] source
• ?Could not verify

  The indexed passages (labeled Abstract; page unavailable) discuss replay during waking immobility and sleep, and report increased maze-related sequences from pre- to postexperience sleep. Searches for replay fidelity and wake-versus-sleep comparisons found no passage saying that sleep replay recapitulates experience less faithfully than wake replay. (LLM, conf=0.76)

  cites [154] sourceExperimental — accuracy not yet measured
• Supported

  The Fidelity of Replay section states that awake sharp-wave ripples contained overall higher-fidelity recapitulations of previously experienced trajectories than ripples recorded in sleep-like states. The Role for ‘Noisy’ Replay likewise describes sleep replay as comparatively more random, supporting the cited comparison. (LLM, conf=0.94)

  cites [155] source

In the paper

“Stimulating somatostatin positive interneurons increased slow-wave amplitudes and their slopes.174”

What the sources say

• Partly supported

  The Results section on chemogenetic activation (page unavailable) reports a 16.8% increase in slow-wave slope after SOM-positive cell activation. It also reports increased slow-wave activity and incidence, but does not establish that the amplitudes of individual slow waves increased; that part of the claim is unsupported. (LLM, conf=0.87)

  cites [174] sourceExperimental — accuracy not yet measured

In the paper

“They also reach the hippocampus possibly via the nucleus reuniens.179”

What the sources say

• Partly supported

  The Introduction (page unavailable) says spindle oscillations are coordinated with hippocampal sharp-wave ripples and that nucleus reuniens connects CA1 and mPFC, making it a candidate to coordinate their activity. The Discussion (page unavailable) also reports spindle–CA1 correlations. These passages support an association and a possible role for reuniens, but do not establish that spindles reach the hippocampus via that route. (LLM, conf=0.82)

  cites [179] sourceExperimental — accuracy not yet measured

In the paper

“While SO-spindle events are traditionally assumed to originate at the cortical level, i.e., from cortical SO driving thalamic spindles, intrathalamic recordings from epilepsy patients suggest additional thalamic contributions, specifically from anterior thalamic nuclei (ANT 188 ).”

What the sources say

• Supported

  The Discussion reports that ANT slow oscillations and spindles preceded their neocortical counterparts and describes the ANT as a hub for coordinating these NREM rhythms. The Results describe SO–spindle complexes recorded in the ANT. No page numbers are provided in the supplied passages. (LLM, conf=0.93)

  cites [188] source

In the paper

“Apart from thalamic afferents and cortical astrocytes, somatostatin positive interneurons inhibiting pyramidal dendrites appear to be specifically implicated in the induction of downstates.161,172,173”

What the sources say

• ?Not checked — no full text of the sourcecites [161] source
• Partly supported

  The Abstract and Results (page not provided) show that cortical astrocytes modulate slow oscillations and that impaired gliotransmission prolongs down-states. The paper does not establish that somatostatin-positive interneurons inhibit pyramidal dendrites or specifically induce downstates; the cited source supports only the astrocyte part of the sentence. (LLM, conf=0.85)

  cites [172] sourceExperimental — accuracy not yet measured
• Partly supported

  The Neocortex section (page unavailable) identifies layer-5 Martinotti cells as cells that intrinsically generate UP and DOWN states, providing limited support for interneuron involvement. It does not identify them as somatostatin-positive, describe inhibition of pyramidal dendrites, or specifically implicate them in inducing DOWN states; the passage instead discusses a current involved in generating their UP states. (LLM, conf=0.82)

  cites [173] sourceExperimental — accuracy not yet measured

In the paper

“In mice, optogenetically reactivating retrosplenial engram ensembles representing a recently acquired contextual fear memory produced a memory trace displaying all features of a remote, systems consolidated memory including higher engagement of neocortical areas during retrieval, contextual generalization, and decreased hippocampal dependence.157”

What the sources say

• Partly supported

  Split into 2 atoms — 0 confirmed, 2 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.

  cites [157] sourceExperimental — accuracy not yet measured

In the paper

“Thus, using a temporospatial association task, Petzka et al. 182 observed in human EEG recordings that spindle amplitudes during postlearning sleep specifically over those areas that had engaged in prior encoding of the task were most distinctly correlated with later retrieval performance.”

What the sources say

• Supported

  The Results section (“EEG results: Spindle amplitude tracks encoding patterns”) reports that greater overlap between post-learning spindle-amplitude topography and encoding topography correlated with better sequence-memory retention from pre- to post-sleep retrieval. It also reports that this relationship was specific to spindle amplitude, rather than spindle duration or density. (LLM, conf=0.91)

  cites [182] source

In the paper

“The anterior thalamus might be specifically involved in coordinating hippocampal memory replay with thalamocortical activity.189”

What the sources say

• Partly supported

  The Abstract and Discussion report that anterodorsal thalamic spindles are temporally coupled with CA1 activity and hippocampal ripples, with stronger coupling after spatial exploration. This supports a possible coordinating role, but the paper measures spiking and ripples, not hippocampal memory replay itself. (LLM, conf=0.84)

  cites [189] sourceExperimental — accuracy not yet measured

In the paper

“Intriguingly, this effect only appeared when stimulation was performed during sleep or light anesthesia, but not during wakefulness.”

What the sources say

• Supported

  The Discussion states that rapid consolidation occurred during sleep and light anesthesia, but not in active awake mice. The Results likewise distinguish stimulation during isoflurane or natural sleep from stimulation during active wakefulness; the wakefulness comparison is specifically for active awake states. (LLM, conf=0.94)

  cites [157] source

In the paper

“This is generally achieved by brain oscillations that, with a shared frequency, synchronize membrane potential depolarization and hyperpolarization across extended neuronal networks.159,160”

What the sources say

• ?Not checked — no full text of the sourcecites [159] source
• Partly supported

  In the Abstract and Conclusions (pages unavailable), Fries describes rhythmic excitation and inhibition coordinated between pre- and postsynaptic groups, producing precisely timed neuronal communication. The paper supports coordination between connected groups, but does not establish synchronized membrane-potential depolarization and hyperpolarization across extended networks generally, or specifically describe distributed neuronal reactivations. (LLM, conf=0.82)

  cites [160] sourceExperimental — accuracy not yet measured

In the paper

“In humans, higher spindle density during sleep after learning of object-word pairs and scene-word pairs increased functional connectivity between hippocampus and neocortex during recall testing on the next day, with this effect pertaining to functional connectivity between anterior hippocampus and ventromedial prefrontal cortex (vmPFC) for object-word pairs, and between posterior hippocampus and”

What the sources say

• Partly supported

  Split into 2 atoms — 1 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.

  cites [34] 34. Cowan, E., Liu, A., Henin, S., Kothare, S., Devinsky, O., and Davachi, L. (2020). Sleep spindles promote the restructuring of memory representa- tions in ventromedial prefrontal cortex through enhanced hippocampal- cortical functional connectivity. J. Neurosci. 40, 1909-1919. https://doi. org/10.1523/JNEUROSCI.1946-19.2020. sourceExperimental — accuracy not yet measured

In the paper

“Indeed, sleep spindles have been found to be associated with signs of increased synchronization between hippocampal and neocortical activity in both mice 189 and humans.34”

What the sources say

• Supported

  The Abstract reports enhanced hippocampal–thalamocortical coupling during spindles in male mice, and the Discussion reports phase-locking of hippocampal ripples to neocortical spindles. This supports the mouse portion attached to this source; the human portion has a separate citation. (LLM, conf=0.88)

  cites [189] source
• Supported

  Citation 34 is attached to the human portion of the sentence; the mice portion has a separate citation. In the Results section on overnight fast spindle density and next-day functional connectivity, the paper reports that fast spindle activity during sleep was associated with increased hippocampal–neocortical functional connectivity measured the next day. That supports the claim’s cautious wording, “signs of increased synchronization,” in humans. (LLM, conf=0.87)

  cites [34] 34. Cowan, E., Liu, A., Henin, S., Kothare, S., Devinsky, O., and Davachi, L. (2020). Sleep spindles promote the restructuring of memory representa- tions in ventromedial prefrontal cortex through enhanced hippocampal- cortical functional connectivity. J. Neurosci. 40, 1909-1919. https://doi. org/10.1523/JNEUROSCI.1946-19.2020. source

In the paper

“In a recent study, ripples induced optogenetically in CA2 neurons following encoding of a social memory task enhanced recall 24 h later.194”

What the sources say

• Supported

  The ‘Triggered CA2 SWRs extend social memory’ section reports that optogenetic stimulation of CA2 pyramidal cells triggered ripples during the delay after social learning. It states that the resulting increase in ripples significantly improved social memory measured 24 hours after learning. (LLM, conf=0.94)

  cites [194] source

In the paper

“However, hippocampal ripples can facilitate cortical SOs.”

What the sources say

• Supported

  The “SOs and ripples” section reports that hippocampal ripples were followed 200–500 ms later by increased cortical EEG SO events, consistent with a bottom-up influence. The Introduction also says ripples might contribute to neocortical SO occurrence. Together, these support the qualified claim that ripples can facilitate cortical SOs. (LLM, conf=0.91)

  cites [199] source
• ?Not checked — no full text of the sourcecites [200] source

In the paper

“Field potentials and unit activity during phasic REMs resemble those observed during image presentation in the wake state.217”

What the sources say

• Supported

  The Results section reports that a late medial temporal lobe field-potential component after REMs resembled the response to image presentation during fixation. It also describes post-REM neuronal firing patterns as reminiscent of responses in the controlled visual experiment. Page numbers were not provided in our copy. (LLM, conf=0.91)

  cites [217] source

In the paper

“At the morphological level, there is evidence that synaptic downscaling during sleep is not ubiquitous but may spare some select synapses that are particularly large or were involved in prior encoding, resulting in increased connectivity at these synapses 163,164,228,231 (Figure 2C).”

What the sources say

• Partly supported

  The Abstract reports that sleep-related downscaling is selective and spares some large synapses. It does not establish that the spared synapses were involved in prior encoding or that their connectivity increased; it says only that a few synapses might upscale during sleep. No page numbers are available in our copy. (LLM, conf=0.88)

  cites [163] sourceExperimental — accuracy not yet measured
• Partly supported

  The paper reports that sleep after motor learning promotes new spine formation on selected dendritic branches in mouse motor cortex, and that sleep improves the survival of some new spines (Abstract; page unavailable). This supports selective increases in learning-related connectivity, but the paper does not show that particularly large synapses are spared or directly test whether synaptic downscaling spares those synapses. (LLM, conf=0.86)

  cites [164] sourceExperimental — accuracy not yet measured
• Partly supported

  The imaging passage (labeled Abstract in our copy; no page given) shows that sleep-related changes were not uniform: some spines gained GluA1, and low-GluA1 spines showed no change in GluA1 or size. But the larger, high-GluA1 spines disproportionately lost GluA1 and shrank. The passage does not establish that synapses involved in prior encoding were spared or that connectivity increased at them. (LLM, conf=0.88)

  cites [228] sourceExperimental — accuracy not yet measured

In the paper

“Hippocampal ripple activity is, in addition, suppressed during the SO downstate.198,199”

What the sources say

• ?Not checked — no full text of the sourcecites [198] source
• Supported

  The “SOs and ripples” section reports suppression of hippocampal ripples around the SO downstate peak in frontal and parietal EEG recordings. The “Triple co-occurrence” section also reports suppressed ripple power around the downstate peak, most distinctly for isolated SOs. (LLM, conf=0.94)

  cites [199] source

In the paper

“Ripples during SWS tend to nest into the excitable troughs of the spindle oscillation thereby forming spindle-ripple events.[195][196][197]”

What the sources say

• ?Not checked — no full text of the sourcecites [195] source
• ?Not checked — no full text of the sourcecites [196] source
• Supported

  The Results (Event-locked analysis; page not provided) report that ripple bursts were nested in individual spindle troughs and describe these troughs as windows of synaptic excitability. The Discussion likewise describes spindle–ripple coupling, and the Results report ripples occurring with spindles during SWS. (LLM, conf=0.94)

  cites [197] source

In the paper

“Likewise, rats training a prehension skill task initially showed increased hippocampal ripples coupled to SOs in primary motor cortex during post-training SWS.201”

What the sources say

• Supported

  The Methods (Behaviour; page unavailable) describe rats learning a reach-to-grasp pellet task. In “Stepwise drop in M1 SO–SWR coupling” (page unavailable), the paper reports significantly greater coupling of hippocampal sharp-wave ripples with primary motor cortex slow oscillations during post-training sleep in early learning (days 1–4); it describes the monitored sleep as non-rapid eye movement sleep. (LLM, conf=0.93)

  cites [201] source

In the paper

“Importantly, this comparison revealed that the max spines-although also undergoing some downscaling-kept a relatively higher GLuA1 expression after sleep than after a post-training period of wakefulness.”

What the sources say

• Supported

  The Results section defines max spines as those with the largest post-learning SEP-GluA1 increase and reports that they lost some SEP-GluA1 after both sleep and sleep deprivation. It says they were in a stronger position *relative to other spines* after sleep, because other spines lost SEP-GluA1 only after sleep; it does not report higher absolute expression in max spines after sleep than after wakefulness. (LLM, conf=0.92)

  cites [165] source

In the paper

“At the level of global network regulation, sleep is considered to primarily serve the downregulation, i.e., in a strict sense a down-selection of synaptic connectivity.223”

What the sources say

• Supported

  In “Heuristic rules for neuronal plasticity in wake and sleep” (page not provided), the paper describes sleep-related synaptic renormalization as competitive down-selection: some synapses become less effective than others. It also explains that less-selected synapses can be depressed and eventually eliminated, supporting the cited characterization of sleep’s role in regulating synaptic connectivity. (LLM, conf=0.91)

  cites [223] source

In the paper

“Using transcranial direct current stimulation and transcranial magnetic stimulation, studies in humans provided functional evidence for sleep decreasing cortical excitability.230”

What the sources say

• Supported

  The Results section (“Sleep deprivation upscales cortical excitability”; page unavailable) reports greater intracortical excitability after sleep deprivation than after sufficient sleep, based on TMS measures. The paper also used tDCS to test sleep-dependent plasticity, with TMS-measured responses. Thus, the claim is supported as a comparison between sufficient sleep and sleep deprivation, rather than a direct measurement of excitability falling during sleep. (LLM, conf=0.87)

  cites [230] source

In the paper

“The primary effect of sleep might well be to reduce overall network activity, and connectivity at excitatory synapses 233 by increasing inhibition, i.e., by pushing the cortical E-I balance toward increased synaptic inhibition.”

What the sources say

• Partly supported

  The Results (Spine shrinkage by two-color glutamate and GABA uncaging; page unavailable) report GABA-associated spine shrinkage, elimination, and reduced glutamate sensitivity, supporting a local mechanism for weakening excitatory connections. The paper does not examine sleep or establish that sleep reduces overall network activity or shifts cortical excitation–inhibition balance toward inhibition. (LLM, conf=0.87)

  cites [233] sourceExperimental — accuracy not yet measured

In the paper

“During succeeding REM sleep epochs activity of these cells decreased, in parallel with the overall population.”

What the sources say

• Supported

  The Results section, under “REM sleep renormalizes calcium activity of spindle-active cells,” states that spindle-active pyramidal cells and the general pyramidal-cell population both decreased their calcium activity during REM sleep. The Discussion likewise describes their REM-sleep dynamics as very similar. No page numbers are provided in the supplied passages. (LLM, conf=0.95)

  cites [166] source

In the paper

“For example, dendrites in layer 1 appear to mainly encode more abstract information compared with deeper layers where perceptual features are encoded.241”

What the sources say

• Supported

  In “Neocortical Layer 1 - The Memory Layer?”, the authors propose that tuft dendrites support learning of context and high-level concepts, while synapses in the other cortical layers support learning of features. The claim’s “appear to” appropriately reflects that this is a hypothesis. (LLM, conf=0.90)

  cites [241] source

In the paper

“with the latter probably particularly contributing to the regulation of synaptic plasticity during development.231,249”

What the sources say

• Supported

  The Discussion (page not supplied) says REM sleep prunes, strengthens, and maintains new synapses during development. It further suggests that activity during REM sleep is perhaps uniquely important for maintaining those synapses, supporting the claim’s qualified emphasis on REM sleep. (LLM, conf=0.93)

  cites [231] source
• Supported

  The Discussion says REM sleep enhances experience-dependent plasticity in the developing cerebral cortex of cats and supports a role for REM sleep in early-life circuit formation. It also describes the results as consistent with a synaptic consolidation role for REM sleep, supporting the citing sentence’s tentative wording. (LLM, conf=0.90)

  cites [249] source

In the paper

“By contrast, REM sleep was found to be characterized by enhanced synaptic pruning that selectively omitted subsets of synapses formed during encoding of new motor memory prior to sleep.231”

What the sources say

• Supported

  The Discussion states that REM sleep immediately after motor learning prunes newly formed spines. The Abstract also describes selective elimination of newly formed synapses after motor learning, matching the claim about pruning a subset of new synapses. (LLM, conf=0.92)

  cites [231] source

In the paper

“Compared with adults, infants and children spent more time in SWS and REM sleep”

What the sources say

• ?Could not verify

  The Abstract says children spend much of their early years asleep, and “Nap Transitions in Infants and Children” describes newborns’ total sleep time, but neither compares time in SWS or REM with adults. Searches for REM sleep and age-related sleep-stage duration found no passage in our copy making that comparison; page numbers were unavailable. (LLM, conf=0.74)

  cites [246] sourceExperimental — accuracy not yet measured
• ?Could not verify

  The Introduction (p. 193) describes a study of sleep EEG in ages 6–18, and EEG Recording and Analysis discusses sleep duration only in explaining the analysis window. Neither passage reports whether infants or children spend more time in SWS and REM sleep than adults; searches for that comparison found no supporting passage. (LLM, conf=0.75)

  cites [248] sourceExperimental — accuracy not yet measured

In the paper

“Thus, similar to spine formation during learning, also spine formation during post-training SWS only pertained to select dendritic branches of motor cortex pyramidal cells.164”

What the sources say

• Partly supported

  The Abstract reports that post-training sleep promotes spine formation on a subset of dendritic branches of mouse motor-cortex layer V pyramidal neurons. The sleep-deprivation and REM-deprivation passages support sleep-dependent, branch-specific formation, but the paper does not establish that the spines formed specifically during slow-wave sleep rather than another part of non-REM sleep. (LLM, conf=0.84)

  cites [164] sourceExperimental — accuracy not yet measured

In the paper

“On a morphological level, SWS following motor task training was found to support branch-specific spine formation in a subset of cells in motor cortex.164”

What the sources say

• Partly supported

  The Abstract reports that sleep after motor learning promotes spine formation on a subset of dendritic branches in mouse motor-cortex neurons, and that task-related neurons reactivate during NREM sleep. It does not specifically identify slow-wave sleep (SWS) as the supporting sleep stage, or describe the spine formation as occurring in a subset of cells rather than a subset of branches. (LLM, conf=0.86)

  cites [164] sourceExperimental — accuracy not yet measured

In the paper

“Corresponding effects were found in humans.234”

What the sources say

• Partly supported

  The Abstract (page unavailable) reports human evidence that cortical excitability depends on sleep homeostasis and describes this study’s measurements of excitation/inhibition balance during sleep deprivation. But the Discussion (page unavailable) says the measured balance returned to baseline without sleep and may be under a prominent circadian influence. The paper therefore supports changes in human excitation/inhibition balance across the sleep–wake cycle, but not an unqualified attribution of those changes to sleep itself. (LLM, conf=0.82)

  cites [234] sourceExperimental — accuracy not yet measured

In the paper

“Moreover, cortical and hippocampal neuronal activity showed more robust decreases across periods of REM sleep than SWS.166,232”

What the sources say

• Partly supported

  The Results (page unavailable) report decreased calcium activity in cortical pyramidal cells during both SWS and REM sleep; the Discussion describes broader cortical downregulation during REM sleep. The paper does not establish that the decreases were larger during REM than SWS, and it did not measure hippocampal neurons; its Discussion only mentions a separate hippocampal study. (LLM, conf=0.86)

  cites [166] sourceExperimental — accuracy not yet measured
• Partly supported

  The Results report that hippocampal firing rates decreased during REM episodes but increased during non-REM episodes. The Discussion says it remains unknown whether these CA1 findings extend to other cortical regions, so the claim’s cortical component is not supported by this paper. (LLM, conf=0.90)

  cites [232] sourceExperimental — accuracy not yet measured

In the paper

“By contrast, the scarce available evidence suggests that such coupling might be weak or even entirely absent during infancy.275”

What the sources say

• Partly supported

  The Results (Hippocampal-Cortical Oscillatory Coupling Through Development) report that significant SO–spindle phase coupling emerged by postnatal day 32 in rats, while the number of temporally coupled events increased with age. This supports weaker coupling at younger ages, but the paper also observes temporal coupling during the study period; it does not establish that coupling is entirely absent during infancy generally. (LLM, conf=0.85)

  cites [275] sourceExperimental — accuracy not yet measured

In the paper

“The greater gain in explicit knowledge in the children was associated with higher amounts of slow oscillatory activity.263”

What the sources say

• ?Could not verify

  The Results report greater slow-wave slopes in prepubertal children than in adolescents, and the introduction mentions memory consolidation generally. Neither passage reports gains in explicit knowledge or tests whether those gains were associated with slow oscillatory activity (searched: explicit knowledge, learning, memory consolidation). (LLM, conf=0.76)

  cites [263] sourceExperimental — accuracy not yet measured

In the paper

“Findings generally agree with the notion that sleep supports processing of episodic-like memories and its precursors also at this early developmental stage”

What the sources say

• Supported

  The Discussion reports converging evidence that sleep supports declarative memory consolidation in the first year of life. The paper identifies deferred imitation as a measure of nonverbal declarative memory and reports that 6- and 12-month-old infants who napped after learning retained the target actions. (LLM, conf=0.86)

  cites [258] source
• Supported

  The Discussion (page unavailable) says sleep close to learning benefits episodic learning in preschool children and that slow-wave sleep during naps may help consolidate those memories. The Introduction also reports evidence of sleep-related memory benefits in young children, supporting the broader wording of the claim. (LLM, conf=0.91)

  cites [267] source

In the paper

“Overall, these studies indicate sleepdependent benefits across a large variety of memory tasks, such as memory for objects and actions, declarative, emotional and language-related memories”

What the sources say

• ?Not checked — no full text of the sourcecites [257] source
• Supported

  The Abstract reports that naps benefited infants’ declarative memory for novel actions and describes earlier sleep benefits in infant language-processing tasks. It also reports benefits for declarative memory in preschool- and school-aged children; the cited child study on emotional declarative memories is identified in the References. Together, these passages support the stated range of tasks across infants and children. (LLM, conf=0.83)

  cites [258] source

In the paper

“and a similar parallel developmental course between the emergence of SOspindle coupling and of object-in-place memory has been shown in rat pups.275”

What the sources say

• Supported

  The Results section reports that significant slow-oscillation–spindle phase coupling appeared by postnatal day 32, which it identifies as the day memory capacities appeared. The paper tested object-place recognition in rat pups and reports that they achieved adult-like performance by that age. Page numbers are not provided in our copy. (LLM, conf=0.93)

  cites [275] source

In the paper

“In even younger 6-to 8-month-old infants, indicators of memory for sequential rules (nonadjacent dependencies in spoken artificial word strings) were found to be enhanced after sleep.271”

What the sources say

• Partly supported

  The Results report indicators of memory for nonadjacent dependencies in both the nap and wake groups of 6- to 8-month-olds. The Discussion says sleep altered a specific stage of processing, but memory of the regularities was generally independent of sleep. Thus, describing the indicators as broadly enhanced after sleep overstates the finding. (LLM, conf=0.86)

  cites [271] sourceExperimental — accuracy not yet measured

In the paper

“Specifically, toddlers who later remembered where, in which temporal order, and in the presence of which toy character they had heard the song exhibited stronger hippocampal activation when the song was played to them during sleep.”

What the sources say

• Partly supported

  The Results (fMRI results; page unavailable) report stronger right-hippocampal activation during sleep in association with better memory for temporal order. They report no such association for spatial location. Although the Methods (Procedure; page unavailable) mention puppets, the paper does not report an activation association with remembering which puppet was present; the Introduction attributes a room-and-puppet finding to an earlier study. (LLM, conf=0.90)

  cites [272] sourceExperimental — accuracy not yet measured
• Partly supported

  The Introduction reports that 28-month-olds showed greater hippocampal activation during sleep for newly learned songs than for novel songs, and that activation was associated with memory for events linked to the songs. It does not specify memory for where or in what order the songs were heard, or which toy character was present. The paper reports this as a finding of earlier studies, rather than its own song experiment. (LLM, conf=0.88)

  cites [273] sourceExperimental — accuracy not yet measured

In the paper

“Children of similar age (7-12 years) showed likewise a greater sleep-dependent gain than adult controls for associative memories between artificial objects and their functions.264”

What the sources say

• Supported

  The Participants section gives the children's age range as 7–12 years and identifies an adult comparison group. The Materials section describes associations between unfamiliar non-objects and their imaginary functions. In Results, children had higher retention indices than adults after sleep, with a significant age-by-condition interaction; the groups did not differ significantly in the wake condition. (LLM, conf=0.94)

  cites [264] source

In the paper

“Sleep, thus, helps the infants to acquire the semantic meaning of words.”

What the sources say

• Supported

  The Discussion states that a short nap preserves specific word meanings and helps infants form generalized word meanings. This supports the claim that sleep helps infants acquire semantic word meanings. (LLM, conf=0.93)

  cites [269] source

In the paper

“For example, testing 7-to 11-year-old children's spatial memories for card pairs showing the same object, Prehn-Kristensen et al. 262 found superior recall after post-encoding nocturnal sleep, in comparison with a daytime wake interval only when these memories were associated with a high reward.”

What the sources say

• Supported

  The Methods describe an object-location task using matching card pairs and a 12-hour interval with overnight sleep or daytime wakefulness. The Results report that, among children, high-rewarded pairs were forgotten less after sleep than after wakefulness, while no other comparison was significant. Page numbers are not available in our copy. (LLM, conf=0.93)

  cites [262] source

In the paper

“There is consistent evidence of an association of sleep spindles with memory consolidation in infancy and childhood (e.g., Friedrich et al. 269 and Friedrich et al. 274 ).”

What the sources say

• Partly supported

  The Discussion reports a correlation between sleep spindle activity and semantic generalization in infants aged 9–16 months, supporting the infancy part of the claim. The paper does not establish consistent evidence across infancy and childhood: it describes its infant result as the first such correlation, while the childhood evidence appears only as a cited preschool study in the References. No page numbers are available in our copy. (LLM, conf=0.84)

  cites [269] sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites [274] source

In the paper

“Friedrich et al. 269 found in 9-to 16-month-old infants that the occurrence of sleep during a 1.5-h napping period after encoding facilitated the generalization of the words to novel but similar objects, indicating a categorization process.”

What the sources say

• Supported

  The Abstract reports that infants aged 9–16 months who napped during the approximately 1.5-hour retention period generalized words to novel category exemplars, whereas infants who stayed awake did not. Methods > Stimuli identifies the spoken words as pseudo-words, and Methods > Experimental design describes the similar objects used to build categories. Page numbers are unavailable in our copy. (LLM, conf=0.92)

  cites [269] source

In the paper

“In humans, the ability to form episodic memories for events bound into a specific spatiotemporal context already emerges at an age between 3 and 4 years.266”

What the sources say

• Partly supported

  The General Discussion (page unavailable) reports that 4-year-olds could bind list contexts with items when the context cue was made more salient. But the study did not test 3-year-olds or establish that this ability emerges between ages 3 and 4; it found that more complex binding continues to develop later. (LLM, conf=0.84)

  cites [266] sourceExperimental — accuracy not yet measured

In the paper

“with some findings suggesting that sleep in infancy particularly supports the formation of abstracted and generalized representations.268”

What the sources say

• Supported

  The Abstract says the studies reviewed show sleep facilitating generalization in infants, in contrast to enhanced precise memory after 18–24 months. The “Sleep and memory in infancy” section likewise describes generalization after sleep alongside poor memory for specific details, supporting the citing sentence’s qualified wording. (LLM, conf=0.93)

  cites [268] source

In the paper

“the same group also demonstrated a facilitating effect of sleep on the formation of episodic-like memory, i.e., the memory for a unique word-object association.270”

What the sources say

• Supported

  The Introduction states that the study tested 14- to 17-month-old infants and found that infants who napped retained individual object–word pairings, whereas those who stayed awake did not. The Results section, “Episodic memories were formed in infants of the nap group,” reports an episodic-context memory response in the nap group that was absent in the wake group. (LLM, conf=0.94)

  cites [270] source

In the paper

“and impaired memory after sleep restriction (e.g., Lo et al. 259 ).”

What the sources say

• Partly supported

  The Methods (Participants; page unavailable) identify the participants as adolescents aged 15–19, not infants or younger children. The Discussion (The impact of sleep in retaining important memories; page unavailable) reports no significant difference in overall memory performance between the sleep-restricted and control groups. The Conclusion (page unavailable) supports the narrower finding that sleep restriction may attenuate the memory benefit of prioritizing information. (LLM, conf=0.86)

  cites [259] sourceExperimental — accuracy not yet measured

In the paper

“Sleep-independent memory formation in starved flies depended on the hunger signal neuropeptide F.”

What the sources say

• Supported

  The passage on NPF signaling reports that starved flies lacking the NPF receptor required sleep for long-term memory, and that knocking down NPF or its receptor likewise made memory sleep-dependent. Extended Data Fig. 5 explicitly describes NPF signaling as essential for sleep-independent memory in starved flies. (LLM, conf=0.94)

  cites [7] 7. Chouhan, N.S., Griffith, L.C., Haynes, P., and Sehgal, A. (2021). Availabil- ity of food determines the need for sleep in memory consolidation. Nature 589, 582-585. https://doi.org/10.1038/s41586-020-2997-y. source

In the paper

“However, the offline replay of ensemble firing patterns in conjunction with hippocampal ripples occurs also during quiet wakefulness.138,150”

What the sources say

• Supported

  The Introduction states that hippocampal replay is associated with sharp-wave ripples during offline periods in both sleep and non-exploratory waking states. It also describes awake replay during pauses in exploration, matching the claim about quiet wakefulness. (LLM, conf=0.94)

  cites [138] source
• Partly supported

  The Results section (“Reactivation of 2D trajectories after random foraging”; page unavailable) reports trajectory reactivation during sharp-wave ripples in quiet immobility after exploration. But the Methods (“Novel Environment Analysis”; page unavailable) calls the rest sessions sleep periods, so this paper does not establish that the recorded offline replay occurred during quiet wakefulness. (LLM, conf=0.83)

  cites [150] sourceExperimental — accuracy not yet measured

In the paper

“Along this line, wake replay is expected not to interfere with consolidation only when it is not ''offline'' but occurs briefly after encoding in the very same context as the encoded event.284,285,293”

What the sources say

• Partly supported

  The Results report that activity during awake sharp-wave ripples promotes stabilization of hippocampal place representations, and the Discussion links this effect to learning new map configurations. The paper does not establish that wake replay avoids interfering with consolidation only if it occurs briefly after encoding in the same context; its manipulation took place during learning, and it makes no such exclusive claim. (LLM, conf=0.85)

  cites [284] sourceExperimental — accuracy not yet measured
• Partly supported

  The Abstract describes awake replay during learning and distinguishes its role from rest/sleep SWR activity associated with consolidation. It does not say that wake replay avoids interfering with consolidation only if it occurs briefly after encoding in the same context; that timing and exclusivity condition is unsupported. (LLM, conf=0.76)

  cites [285] sourceExperimental — accuracy not yet measured
• Partly supported

  The Introduction (page unavailable) describes repeated encoding followed by free recall, and the Discussion says rehearsal rapidly initiates systems consolidation. But the paper does not say that wake replay avoids interfering with consolidation only when it occurs shortly after encoding in the same context, or establish that contrast with offline replay. (LLM, conf=0.79)

  cites [293] sourceExperimental — accuracy not yet measured

In the paper

“Thus, spontaneous offline replay activity in hippocampal networks enhancing memory indeed appears to be a unique feature of sleep-dependent consolidation 66,74”

What the sources say

• ?Not checked — no full text of the sourcecites [66] 66. Sawangjit, A., Oyanedel, C.N., Niethard, N., Salazar, C., Born, J., and In- ostroza, M. (2018). The hippocampus is crucial for forming non-hippo- campal long-term memory during sleep. Nature 564, 109-113. https:// doi.org/10.1038/s41586-018-0716-8. source
• Partly supported

  The Introduction describes hippocampal replay during sleep and says hippocampal–cortical dialogue is thought to facilitate memory consolidation. The Discussion also proposes that replay during sleep may benefit motor-sequence consolidation. Neither passage establishes that memory-enhancing offline replay is unique to sleep-dependent consolidation. (LLM, conf=0.84)

  cites [74] 74. Schapiro, A.C., Reid, A.G., Morgan, A., Manoach, D.S., Verfaellie, M., and Stickgold, R. (2019). The hippocampus is necessary for the consol- idation of a task that does not require the hippocampus for initial learning. Hippocampus 29, 1091-1100. https://doi.org/10.1002/hipo.23101. sourceExperimental — accuracy not yet measured

In the paper

“Interestingly, in the experiments natural awake consolidation turned out to be also superior to sleep consolidation when object recognition after 1 week was tested in an environmental context entirely different from that during encoding.”

What the sources say

• Supported

  The Discussion (page unavailable) states that object recognition after wake-associated consolidation was superior to that after sleep when rats were tested in a context different from encoding. Materials and Methods (page unavailable) specifies that this context-change test occurred at the 1-week retention interval. (LLM, conf=0.94)

  cites [6] 6. Sawangjit, A., Harkotte, M., Oyanedel, C.N., Niethard, N., Born, J., and Inostroza, M. (2022). Two distinct ways to form long-term object recog- nition memory during sleep and wakefulness. Proc. Natl. Acad. Sci. USA 119. e2203165119. https://doi.org/10.1073/pnas.2203165119. source

In the paper

“Beyond pointing to distinct mechanisms mediating wake and sleep-dependent consolidation processes, the findings show that unlike sleep consolidation, wake consolidation strongly depends on internal organismic states such as hunger.”

What the sources say

• Partly supported

  The Abstract reports that hunger-related NPF signaling helps select the sleep-independent memory pathway in starved flies, supporting the point about wake consolidation. But the Abstract and Discussion describe a feeding/hunger-dependent switch between both pathways; they do not support the contrast that sleep-dependent consolidation is unlike wake consolidation in this respect. The study concerns appetitive memory in Drosophila. (LLM, conf=0.86)

  cites [7] 7. Chouhan, N.S., Griffith, L.C., Haynes, P., and Sehgal, A. (2021). Availabil- ity of food determines the need for sleep in memory consolidation. Nature 589, 582-585. https://doi.org/10.1038/s41586-020-2997-y. sourceExperimental — accuracy not yet measured

In the paper

“Against the backdrop that the mammalian hippocampus shows some homologies with the insect mushroom bodies in structure and function, 283 it is interesting that sleep and wake-associated consolidation has been also dissociated in Drosophila.”

What the sources say

• Supported

  The Abstract reports distinct sleep-dependent and sleep-independent memory consolidation in Drosophila: fed flies needed increased sleep after conditioning, while starved flies did not. It also describes starvation as increasing waking foraging activity at the expense of sleep. (LLM, conf=0.92)

  cites [7] 7. Chouhan, N.S., Griffith, L.C., Haynes, P., and Sehgal, A. (2021). Availabil- ity of food determines the need for sleep in memory consolidation. Nature 589, 582-585. https://doi.org/10.1038/s41586-020-2997-y. source

In the paper

“However, challenging the view of systems consolidation as a process exclusively linked to sleep, signs of systems consolidation can be also observed under certain conditions in the wake state.[290][291][292][293]”

What the sources say

• ?Could not verify

  The Introduction (page unavailable) describes memory reactivation and consolidation during sleep, and the Discussion (page unavailable) addresses spindle timing during sleep. A References entry names a study of reactivation during wakefulness and sleep, but our copy does not state that signs of systems consolidation occur in the wake state. Searches for wakefulness and systems consolidation found no passage establishing that claim. (LLM, conf=0.73)

  cites [290] sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites [291] source
• Supported

  The Discussion says that neuronal reactivation associated with systems consolidation occurs during resting wakefulness as well as sleep. It also reports that repeated encounters with learning material produced signs of systems consolidation, supporting the claim that sleep is not exclusively required under certain conditions. (LLM, conf=0.90)

  cites [292] source
• Supported

  The Discussion states that repeated study is sufficient to trigger systems consolidation, although sleep is needed to stabilize the resulting transitions. This supports the claim that signs of systems consolidation can occur during wakeful rehearsal, rather than exclusively during sleep. (LLM, conf=0.94)

  cites [293] source

In the paper

“By contrast, when starved the flies formed such long-term memory also in sleep-deprived conditions.”

What the sources say

• Supported

  The paper’s main text reports that flies kept starved after training showed no loss of long-term memory following sleep deprivation. Extended Data Fig. 2(e) likewise states that six hours of sleep deprivation had no effect on their long-term memory. (LLM, conf=0.94)

  cites [7] 7. Chouhan, N.S., Griffith, L.C., Haynes, P., and Sehgal, A. (2021). Availabil- ity of food determines the need for sleep in memory consolidation. Nature 589, 582-585. https://doi.org/10.1038/s41586-020-2997-y. source

In the paper

“underlining the importance of naturally low glucocorticoid concentrations for consolidation during nocturnal sleep.”

What the sources say

• Supported

  The Discussion (page unavailable) reports that elevated glucocorticoids impair declarative memory formation during early slow-wave sleep, supporting the importance of keeping levels low for sleep-related consolidation. The paper reports this sleep finding from prior research; its own experiments used mouse hippocampal slices. (LLM, conf=0.84)

  cites [281] source

In the paper

“(D)Synaptic AMPA receptor density increases during learning and is globally reduced during subsequent sleep.”

What the sources say

• Supported

  The Results section, “Learning leads to a net increase in SEP-GluA1 expression,” reports a significant increase in mean spine SEP-GluA1 after motor learning. The Discussion identifies this as increased spine-surface GluA1-containing AMPA receptors in the mouse primary motor cortex; it does not establish an increase in every synapse or brain region. (LLM, conf=0.90)

  cites [165] source

In the paper

“Indeed, conditions such as the globally low levels of acetylcholine and glucocorticoids together with the occurrence of SO, spindles and ripples that hallmark SWS, facilitate hippocampal replay, output, and the transmission of reactivated memory information toward neocortical long-term storage sites.281,[286][287][288][289]”

What the sources say

• Partly supported

  The Results report that high-dose corticosterone reduced sharp-wave frequency in mouse hippocampal slices, and the Discussion links elevated glucocorticoids to impaired memory formation during slow-wave sleep. This supports a role for glucocorticoid levels in memory-related oscillations, but the paper does not establish that the combined conditions of low acetylcholine, slow oscillations, spindles and ripples facilitate replay or transfer of memories to neocortex. Page numbers are unavailable in our copy. (LLM, conf=0.84)

  cites [281] sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites [286] source
• ?Not checked — no full text of the sourcecites [287] source
• Partly supported

  Split into 2 atoms — 0 confirmed, 2 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.

  cites [288] sourceExperimental — accuracy not yet measured
• ?Not checked — no full text of the sourcecites [289] source

In the paper

“(D)Synaptic AMPA receptor density increases during learning and is globally reduced during subsequent sleep.”

What the sources say

• Partly supported

  The Discussion reports a net decrease in spine-surface GluA1-containing AMPA receptors after sleep in adult mouse primary motor cortex. But it explicitly describes sleep-related synaptic decreases as widespread, not global, and says it is unknown whether the result extends to other adult brain areas or layers. The claim overstates both the extent of the reduction and the receptor types measured. (LLM, conf=0.90)

  cites [165] sourceExperimental — accuracy not yet measured

In the paper

“(E) Sleep, particularly REM sleep, globally downregulates calcium activity in cortical pyramidal cells.”

What the sources say

• Supported

  The Results section states that pyramidal-cell calcium activity decreased on average during both SWS and REM sleep. The Abstract reports a general downregulation during REM sleep, including in cells that were active during SWS spindles. Here, “globally” describes the population-level trend, not an identical response in every cell. (LLM, conf=0.93)

  cites [166] source
• ?Not checked — no full text of the sourcecites [167] source

In the paper

“Downscaling includes the synapses that were upscaled during learning, although to a lesser extent.165”

What the sources say

• Partly supported

  The Results report that spines with the largest post-learning GluA1 increases lost some GluA1 over the next seven hours, while other spines lost GluA1 after sleep. But the learning-associated spines also lost GluA1 after sleep deprivation, so the paper does not establish that sleep-dependent downscaling included them; the Discussion also notes that regression to the mean may explain some of their decrease (pages not provided). (LLM, conf=0.86)

  cites [165] sourceExperimental — accuracy not yet measured

In the paper

“Cells that are particularly active during sleep spindles are spared from this downregulation during non-REM sleep where these cells even increase their activity.166,167”

What the sources say

• Supported

  The Discussion (Figure 6 summary) states that spindle-active pyramidal cells increase their activity during slow-wave sleep while the remaining pyramidal cells decrease theirs. This supports the claim that these spindle-active cells are spared from the usual downregulation during this phase of non-REM sleep. (LLM, conf=0.92)

  cites [166] source
• ?Not checked — no full text of the sourcecites [167] source

In the paper

“Selective optogenetic suppression of hippocampal theta activity during REM sleep impaired consolidation of object-location memory.221”

What the sources say

• ?Not checked — no full text of the sourcecites [221] source

In the paper

“Interestingly, ripples appear to be also directly implicated in synaptic scaling, as closed-loop optogenetic suppression of hippocampal ripples in dorsal CA3 prevented spontaneous synaptic downscaling in these circuits.132”

What the sources say

• ?Not checked — no full text of the sourcecites [132] 132. Contreras, M.P., Fechner, J., Born, J., and Inostroza, M. (2023). Acceler- ating maturation of spatial memory systems by experience -evidence from sleep oscillation signatures of memory processing. J. Neurosci. JN-RM-1967-22. https://doi.org/10.1523/JNEUROSCI.1967-22.2023. 279. Kelemen, E., Bahrendt, M., Born, J., and Inostroza, M. (2014). Hippocam- pal corticosterone impairs memory consolidation during sleep but im- proves consolidation in the wake state. Hippocampus 24, 510-515. https://doi.org/10.1002/hipo.22266. 280. Wilhelm, I., Wagner, U., and Born, J. (2011b). Opposite effects of c source

In the paper

“In specific task conditions, this benefit can be even greater than in the mature brain”

What the sources say

• ?Not checked — no full text of the sourcecites [265] source

In the paper

“Interestingly, in rat pups the sleep-dependent occurrence of adult-like persisting spatial object-in-place representations could be advanced to PD31 (corresponding to late childhood) by repeatedly exposing them to 5-min periods of discrete spatial experiences (configurational changes in two identical objects) on the days before 278”

What the sources say

• ?Not checked — no full text of the sourcecites [278] source

In the paper

“Contrasting with this hypothesis, children around 10 years of age showed greater sleep-dependent gains in explicit knowledge of a sequence than their adult controls.46”

What the sources say

• ?Not checked — no full text of the sourcecites [46] 46. Wilhelm, I., Rose, M., Imhof, K.I., Rasch, B., B€ uchel, C., and Born, J. (2013). The sleeping child out-plays the adult's capacity to convert im- plicit into explicit knowledge. Nat. Neurosci. 16, 391-393. https://doi. org/10.1038/nn.3343. source

In the paper

“Moreover, in school children aged 7-15 years the coupling strength of spindles to the SO upstate during post-encoding sleep predicted recall of word lists on the next day, with this relationship occurring independently of the children's age, 277”

What the sources say

• ?Not checked — no full text of the sourcecites [277] source

In the paper

“some studies in children suggest more robust benefits from sleep for episodic and declarative types of memory in comparison with task aspects less reliant on the hippocampus, similar to procedural motor skills or object recognition.257,260”

What the sources say

• ?Not checked — no full text of the sourcecites [257] source
• ?Not checked — no full text of the sourcecites [260] source

In the paper

“For example, 6-month-old infants following a nap showed signs of memory for semantic categories associated to artificial words, which were positively correlated with local spindle activity during the nap.274”

What the sources say

• ?Not checked — no full text of the sourcecites [274] source

In the paper

“Inhibiting the hippocampus by muscimol injection during post-encoding sleep impaired object recognition tested 1 week later.”

What the sources say

• ?Not checked — no full text of the sourcecites [66] 66. Sawangjit, A., Oyanedel, C.N., Niethard, N., Salazar, C., Born, J., and In- ostroza, M. (2018). The hippocampus is crucial for forming non-hippo- campal long-term memory during sleep. Nature 564, 109-113. https:// doi.org/10.1038/s41586-018-0716-8. source

In the paper

“Although long-term memory consolidation is supported by sleep, it is unclear how it differs from that during wakefulness.”

• Also stated in: § INTRODUCTION · boosted

  “Sleep has been known to support the formation of long-term memory since the first experimental evidence was provided about a century ago.”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Rasch and Born, 3
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Klinzing et al., 4
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Girardeau et al. 5

In the paper

“hundreds of studies in humans and animals have shown that in comparison with wakefulness, sleep following the encoding of experimental stimuli, produces more long-lasting and stable memories”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Rasch and Born, 3
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Klinzing et al., 4
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Girardeau et al. 5

In the paper

“it also explains sleep benefits for non-hippocampus-dependent kinds of memory (e.g., procedural skills), as such memories are naturally experienced in the context of specific episodes implicating that they are always embedded into hippocampus-dependent episodic representations.”

In the paper

“Across a great variety of tasks, these studies revealed stronger influences of SWS than REM sleep on memory consolidation.”

In the paper

“in mice, the optogenetic induction of spindles has confirmed their causal contribution to memory consolidation.”

What the sources say

• ?Not checked — no full text of the sourcecites [58] 58. Latchoumane, C.V., Ngo, H.V., Born, J., and Shin, H.S. (2017). Thalamic spindles promote memory formation during sleep through triple phase- locking of cortical, thalamic, and hippocampal rhythms. Neuron 95, 424-435.e6. https://doi.org/10.1016/j.neuron.2017.06.025. source

In the paper

“using visual patterns, Lutz et al. 41 found that post-encoding sleep enhanced generalization of memory recall to a category prototype not presented during encoding, at a recall test 1 year but not 1 day after encoding.”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Lutz et al. 41

In the paper

“The link between sleep spindle activity during post-encoding sleep and later memory performance is even more consistent than that for measures of SWA”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Alger et al., 53
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Peyrache and Seibt, 54
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Lutz et al. 55

In the paper

“No robust associations have been found between the EEG signatures of REM sleep and memory consolidation in humans”

In the paper

“place cells coding for specific locations in an environment replay in the same sequential order during sleep as during prior awake spatial navigation.”

What the sources say

• ?Not checked — no full text of the sourcecites [78] 78. Lee, A.K., and Wilson, M.A. (2002). Memory of sequential experience in the hippocampus during slow wave sleep. Neuron 36, 1183-1194. https://doi.org/10.1016/s0896-6273(02)01096-6. source
• ?Not checked — no full text of the sourcecites [79] 79. Wilson, M.A., and McNaughton, B.L. (1994). Reactivation of hippocam- pal ensemble memories during sleep. Science 265, 676-679. https:// doi.org/10.1126/science.8036517. source

In the paper

“The coupling of hippocampal replay to the depolarizing upstates of SOs emerging from neocortical networks and to thalamic spindles represents a mechanism that supports the transmission of the reactivated memory information and its storage, through synaptic consolidation mechanisms, into respective neocortical networks serving as long-term store.”

In the paper

“At the core of this process is the neuronal replay of memory, i.e., repeated reactivations of firing patterns in neuron ensembles that were activated and used for encoding of the episodes during prior wake phases, and which re-occur in hippocampal networks during post-encoding SWS in coordination with hippocampal sharp wave-ripples (SPW-Rs).”

What the sources say

• ?Not checked — no full text of the sourcecites [4] 4. Klinzing, J.G., Niethard, N., and Born, J. (2019). Mechanisms of systems memory consolidation during sleep. Nat. Neurosci. 22, 1598-1610. https://doi.org/10.1038/s41593-019-0467-3. source
• ?Not checked — no full text of the sourcecites [73] 73. Diekelmann, S., and Born, J. (2010). The memory function of sleep. Nat. Rev. Neurosci. 11, 114-126. https://doi.org/10.1038/nrn2762. source

In the paper

“refining this approach by disrupting only specific reactivation events of a prior learning experience (through online decoding of the content of ensemble firing patterns) revealed a selective recall deficit that was restricted to the spatial environment whose reactivation was disturbed.”

What the sources say

• ?Not checked — no full text of the sourcecites [86] 86. Gridchyn, I., Schoenenberger, P., O'Neill, J., and Csicsvari, J. (2020). As- sembly-specific disruption of hippocampal replay leads to selective memory deficit. Neuron 106, 291-300.e6. e296. https://doi.org/10. 1016/j.neuron.2020.01.021. source

In the paper

“Collectively, these findings support the notion that neuronal reactivations of newly encoded experiences during sleep specifically contribute to the reorganization of respective representations in hippocampal-cortical systems.”

In the paper

“Aly et al. 97 showed that the conjoint offline reactivation of engram cells in anterior cingulate cortex (ACC) coding for distinct experiences is necessary for forming generalized representations of spatial context during sleep.”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Aly et al. 97

In the paper

“using representational similarity analysis, Cairney et al. 117 reliably identified the image category (object vs. scene) associated with the verbal cue (adjectives) in the raw EEG traces during a time window of increased fast spindle power 1.7-2.3s after cue onset.”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Cairney et al. 117

In the paper

“several studies succeeded in identifying signs of spontaneous memory reprocessing during post-learning sleep”

What the sources say

• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Rubin et al., 110
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Sterpenich et al., 111
• ?Not matched

  Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  cites Zhang et al. 112

In the paper

“They also share the factors that select which content is preferably reactivated, due to saliency or relevance, including novelty and reward.”

What the sources say

• ?Not checked — no full text of the sourcecites [80] 80. Diba, K., and Buzsa ´ki, G. (2007). Forward and reverse hippocampal place-cell sequences during ripples. Nat. Neurosci. 10, 1241-1242. https://doi.org/10.1038/nn1961. source
• ?Not checked — no full text of the sourcecites [81] 81. Foster, D.J., and Wilson, M.A. (2006). Reverse replay of behavioural se- quences in hippocampal place cells during the awake state. Nature 440, 680-683. https://doi.org/10.1038/nature04587. source
• ?Not checked — no full text of the sourcecites [141] source

In the paper

“the experience of an entirely novel environment inducing the emergence of a new hippocampal map produced sustained reactivations during sleep for up to 7 h with a peak after 3-5 h.”

What the sources say

• ?Not checked — no full text of the sourcecites [133] 133. Antony, J.W., Piloto, L., Wang, M., Pacheco, P., Norman, K.A., and Paller, K.A. (2018). Sleep spindle refractoriness segregates periods of memory reactivation. Curr. Biol. 28, 1736-1743.e4. https://doi.org/10.1016/j. cub.2018.04.020. 291. source

In the paper

“The time course of reactivations might also depend on the familiarity of the encoded environments.”

In the paper

“this temporal dynamic likely represents a general feature of reactivations showing maximum rates within 30 min after encoding also during wakefulness.”

What the sources say

• ?Not checked — no full text of the sourcecites [81] 81. Foster, D.J., and Wilson, M.A. (2006). Reverse replay of behavioural se- quences in hippocampal place cells during the awake state. Nature 440, 680-683. https://doi.org/10.1038/nature04587. source

In the paper

“Assuming that the underlying neuron ensembles are synaptically connected, this dynamic basically argues against an immediate effect of replay ''strengthening'' synaptic connectivity in these ensembles.”

What the sources say

• ?Not checked — no full text of the sourcecites [132] 132. Contreras, M.P., Fechner, J., Born, J., and Inostroza, M. (2023). Acceler- ating maturation of spatial memory systems by experience -evidence from sleep oscillation signatures of memory processing. J. Neurosci. JN-RM-1967-22. https://doi.org/10.1523/JNEUROSCI.1967-22.2023. 279. Kelemen, E., Bahrendt, M., Born, J., and Inostroza, M. (2014). Hippocam- pal corticosterone impairs memory consolidation during sleep but im- proves consolidation in the wake state. Hippocampus 24, 510-515. https://doi.org/10.1002/hipo.22266. 280. Wilhelm, I., Wagner, U., and Born, J. (2011b). Opposite effects of c source

In the paper

“However, in order to consolidate memory representations that are widely distributed across different brain regions, neuronal reactivations need to be temporally precisely synchronized between these distributed areas.”

In the paper

“By phase-coupling local gamma band activities, spindles might serve to synchronize memory processing in distributed cortical networks.”

In the paper

“This contribution might not be conveyed by the replay phenomenon itself but owed to the unique milieu during sleep.”

In the paper

“Altogether, these similarities between reactivations occurring during sleep and wakefulness exclude that reactivation, and replay in the narrow sense, itself is the critical factor that explains superior consolidation effects during sleep.”

In the paper

“Spindles are waxing and waning 11-16 Hz oscillatory events of 0.5-3 s duration that are generated in GABAergic feedback-loops including the nucleus reticularis of the thalamus and via afferent fibers widely spread to the cortex, reaching predominantly central and parietal areas.59”

What the sources say

• ?Not checked — no full text of the sourcecites [59] 59. Fernandez, L.M.J., and L€ uthi, A. (2020). Sleep spindles: mechanisms and functions. Physiol. Rev. 100, 805-868. https://doi.org/10.1152/physrev. 00042.2018. source

In the paper

“Specifically, the embedding of replay within SPWs, spindles, and SOs may enable the large-scale interaction between replay across hippocampal and thalamocortical networks supporting systems consolidation during sleep.”

In the paper

“In this way, oscillations prime synaptic plasticity by timing the firing properties of activated neurons to the excitable depolarizing phase of network oscillations.”