Durrant, Simon

Johnson, Jennifer

Frontiers in Psychology

Jennifer M. Johnson

Simon J. Durrant

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GENERAL COMMENTARYpublished: 19 June 2018doi: 10.3389/fpsyg.2018.01022Frontiers in Psychology | www.frontiersin.org 1 June 2018 | Volume 9 | Article 1022Edited by:Peter Klaver,University of Surrey, United KingdomReviewed by:Emma L. Axelsson,Uppsala University, SwedenJenny Richmond,University of New South Wales,Australia*Correspondence:Simon J. DurrantSiDurrant@lincoln.ac.ukSpecialty section:This article was submitted toDevelopmental Psychology,a section of the journalFrontiers in PsychologyReceived: 08 December 2017Accepted: 31 May 2018Published: 19 June 2018Citation:Johnson JM and Durrant SJ (2018)Commentary: Effects of Sleep onWord Pair Memory inChildren–Separating Item and SourceMemory Aspects.Front. Psychol. 9:1022.doi: 10.3389/fpsyg.2018.01022Commentary: Effects of Sleep onWord Pair Memory inChildren–Separating Item and SourceMemory AspectsJennifer M. Johnson and Simon J. Durrant*School of Psychology, University of Lincoln, Lincoln, United KingdomKeywords: declarative memory consolidation, sleep, children, hippocampus, prefrontal cortexA commentary onEffects of Sleep on Word Pair Memory in Children–Separating Item and Source MemoryAspectsby Wang, J. Y., Weber, F. D., Zinke, K., Noack, H., and Born, J. (2017). Front. Psychol. 8:1533.doi: 10.3389/fpsyg.2017.01533Evidence of sleep-dependent memory consolidation in adults has become widespread in the lasttwo decades (Rasch and Born, 2013). However, investigations of similar consolidation in childrenare still at an early stage (Maski et al., 2015).Previous research has shown sleep-dependent memory consolidation in children on taskssuch as lexical integration of novel words (Henderson et al., 2012) and spatial location learning(Kurdziel et al., 2013). However, this appears to be limited to declarative memory tasks; procedurallearning tasks do not show the same sleep-related benefit in children that is seen in adults (Fischeret al., 2007; Wilhelm et al., 2008). Declarative memory tasks are strongly hippocampal-dependent(Squire, 1992), and consolidation of hippocampal memories is particularly associated with slowwave sleep (SWS) (Diekelmann et al., 2009; Born and Wilhelm, 2012). It has therefore beensuggested that the large amount of SWS seen during childhood (Ohayon et al., 2004) is the reasondeclarative memories show sleep-dependent consolidation in children.However, not all declarative memories are equal. In particular, a key distinction can be drawnbetween the memory of an object (item memory) and the context in which it was encountered(source memory). The recent paper byWang et al. (2017) used a clever design to separate these twoforms of memory, and examined their consolidation in children across an 11 h interval containingwake or sleep or a shorter 1 h wake interval, and compared this to adults with an 11 h sleep or1 h wake interval. The design used two lists of word pairs and the innovation was to measurememory not only for the target words using cued recall (item memory), but also for memory ofwhich list they came from (source memory). When comparing adults to children, a key finding wasthat memory for word pairs was lower after sleep in children (compared to before), but showed noreduction after sleep in adults (Figure 2B in the original paper). Interestingly, this turned out to bedriven entirely by the word pairs without source memory; where the item and source had both beenremembered, no difference was seen.The authors interpreted their findings in the context of less distinct encoding in childrenleading to more rapid unbinding of items and sources. However, this does not explain whyitem-only memory declines across sleep in children but item-source memory remains intact, andwhy the opposite pattern (stronger consolidation across sleep of item-only memory compared toitem-source memory) is seen in adults.Johnson and Durrant Effects of Sleep on Word MemoryWe believe that the influential model by Preston andEichenbaum (2013) allows a more comprehensive frameworkin which to explain the findings. This model postulates theexistence of a “what” stream of cortical processing fromthe dorsal hippocampus through the perirhinal cortex, anda “where” stream from the dorsal hippocampus through theparahippocampal cortex. Additionally, the ventral hippocampusconnects to the medial prefrontal cortex (mPFC), where pre-existing schemata influence subsequent memory processing (Tseet al., 2007; van Kesteren et al., 2010, 2012), and exert a top-down influence on “what” facilitating segregation from “where.”Importantly, Eichenbaum (2014, 2017) has suggested that the“where” stream also processes temporal context information ina similar way, i.e., it may also operate as a “when” stream.A key difference in the declarativememory systems of childrenand adults is that the density of hippocampal synapses in childrenreaches similar levels to adults within the first 6 months oflife (Seress and Ábrahám, 2008), but the frontal lobes havea prolonged developmental course (Luna and Sweeney, 2004)with myelination (Giedd et al., 1999), synaptic density counts(Huttenlocher, 1990) and overall function of the prefrontalcortex not reaching adult levels until late adolescence (Huberand Born, 2014). In children we should therefore expectto see stronger bottom-up processing, and a reduced top-down influence of existing schemata, which would otherwisetypically strengthen item (“what”) memory at the expense of listcontext (“where/when”). This is supported by previous evidenceconfirming that more rapid consolidation of schematic memoriesoccurs during sleep and involves the mPFC (Durrant et al., 2015),and that sleep-dependent consolidation leads to a reduction inassociated contextual information (Cairney et al., 2011).For the Wang et al. (2017) study, greater decontextualizationand consolidation during sleep in adults compared to childrenwould show up as a bigger reduction in item-only memory inchildren and this is exactly what the authors found (Figure 2D).However, we would expect to see relatively little change acrosssleep for items with their original contexts intact in both childrenand adults, reflecting equal hippocampal development, and wesee this too (Figure 2C). This argument is further strengthenedby the fact that the relationship between SWS and memoryconsolidation seen in the study in children was limited to itemswhich retained their context. It also explains why the childrenshowed little sleep-dependent benefit for items which have losttheir contextual cues (Figure 1F).In children, memory for the lists themselves was alsostronger after sleep compared to wake (Figure 1G), while inadults, no significant enhancement of list memory was seen(Supplementary Material). If memory for the lists makes use ofthe contextual stream, the absence of strong mPFC influenceallows both streams to consolidate during sleep, whereas thetop-down mPFC influence in adults strengthens the “what”stream at the expense of the “where/when” stream (hence listmemory is not enhanced after sleep compared to wake inadults).The neural basis of our interpretation could be tested in fMRI,where we predict that prefrontal activation will be stronger inadults during sleep and associated with item-only memory, whilehippocampal activation would be associated with both item andsource memory, but more strongly for items which retain theircontext.The Wang et al. (2017) paper has shown the clear benefitsof measuring different aspects of declarative memory in a singlestudy. We hope this new interpretation will help further improveour understanding of different memory systems and stimulateresearch in this area.AUTHOR CONTRIBUTIONSAll authors listed have made a substantial, direct and intellectualcontribution to the work, and approved it for publication.REFERENCESBorn, J., and Wilhelm, I. (2012). System consolidation of memory during sleep.Psychol. Res. 76, 192–203. doi: 10.1007/s00426-011-0335-6Cairney, S. A., Durrant, S. J., Musgrove, H., and Lewis, P. A. (2011). Sleep andenvironmental context: interactive effects for memory. Exp. Brain Res. 214,83–92. doi: 10.1007/s00221-011-2808-7Diekelmann, S., Wilhelm, I., and Born, J. (2009). The whats and whensof sleep-dependent memory consolidation. Sleep Med. Rev. 13, 309–321.doi: 10.1016/j.smrv.2008.08.002Durrant, S. J., Cairney, S. A., McDermott, C., and Lewis, P. A. (2015). Schema-conformant memories are preferentially consolidated during REM sleep.Neurobiol. Learn. Mem. 122, 41–50. doi: 10.1016/j.nlm.2015.02.011Eichenbaum, H. (2014). Time cells in the hippocampus: A new dimension formapping memories. Nat. Rev. Neurosci. 15, 732–744. doi: 10.1038/nrn3827Eichenbaum, H. (2017). Time (and space) in the hippocampus. Curr. Opin. Behav.Sci. 17, 65–70. doi: 10.1016/j.cobeha.2017.06.010Fischer, S., Wilhelm, I., and Born, J. (2007). Developmental differences in sleep’srole for implicit off-line learning: comparing children with adults. J. Cogn.Neurosci. 19, 214–227. doi: 10.1162/jocn.2007.19.2.214Giedd, J. N., Blumenthal, J., Jeffries, N. O., Castellanos, F. X., Liu, H., Zijdenbos,A., et al. (1999). Brain development during childhood and adolescence: alongitudinal MRI study. Nat. Neurosci. 2, 861–863. doi: 10.1038/13158Henderson, L. M., Weighall, A. R., Brown, H., and Gareth Gaskell, M. (2012).Consolidation of vocabulary is associated with sleep in children. Dev. Sci. 15,674–687. doi: 10.1111/j.1467-7687.2012.01172.xHuber, R., and Born, J. (2014). Sleep, synaptic connectivity, and hippocampalmemory during early development. Trends Cogn. Sci. 18, 141–152.doi: 10.1016/j.tics.2013.12.005Huttenlocher, P. R. (1990). Morphometric study of humancerebral cortex development. Neuropsychologia 28, 517–527.doi: 10.1016/0028-3932(90)90031-IKurdziel, L., Duclos, K., and Spencer, R. M. (2013). Sleep spindles in middaynaps enhance learning in preschool children. Proc. Natl. Acad. Sci. U.S.A. 110,17267–17272. doi: 10.1073/pnas.1306418110Luna, B., and Sweeney, J. A. (2004). The emergence of collaborative brain function:fMRI studies of the development of response inhibition. Ann. N. Y. Acad. Sci.1021, 296–309. doi: 10.1196/annals.1308.035Maski, K., Holbrook, H., Manoach, D., Hanson, E., Kapur, K., and Stickgold,R. (2015). Sleep dependent memory consolidation in children withautism spectrum disorder. Sleep 38, 1955–1963. doi: 10.5665/sleep.5248Ohayon, M. M., Carskadon, M. A., Guilleminault, C., and Vitiello, M. V. (2004).Meta-analysis of quantitative sleep parameters from childhood to old agein healthy individuals: developing normative sleep values across the humanlifespan. Sleep 27, 1255–1273. doi: 10.1093/sleep/27.7.1255Frontiers in Psychology | www.frontiersin.org 2 June 2018 | Volume 9 | Article 1022Johnson and Durrant Effects of Sleep on Word MemoryPreston, A. R., and Eichenbaum, H. (2013). Interplay of hippocampusand prefrontal cortex in memory. Curr. Biol. 23, R764–R773.doi: 10.1016/j.cub.2013.05.041Rasch, B., and Born, J. (2013). About sleep’s role in memory. Physiol. Rev. 93,681–766. doi: 10.1152/physrev.00032.2012Seress, L., and Ábrahám, H. (2008). “Pre-and postnatal morphologicaldevelopment of the human hippocampal formation,” in Handbook ofDevelopmental Cognitive Neuroscience, 2nd Edn, eds C. A. Nelson and M.Luciana (Cambridge, MA: MIT Press), 187–209.Squire, L. R. (1992). Declarative and nondeclarative memory: multiple brainsystems supporting learning and memory. J. Cogn. Neurosci. 4, 232–243.doi: 10.1162/jocn.1992.4.3.232Tse, D., Langston, R. F., Kakeyama, M., Bethus, I., Spooner, P. A., Wood, E.R., et al. (2007). Schemas and memory consolidation. Science 316, 76–82.doi: 10.1126/science.1135935van Kesteren, M. T., Fernández, G., Norris, D. G., and Hermans, E.J. (2010). Persistent schema-dependent hippocampal-neocorticalconnectivity during memory encoding and postencoding rest in humans.Proc. Natl. Acad. Sci. U.S.A. 107, 7550–7555. doi: 10.1073/pnas.0914892107van Kesteren, M. T., Ruiter, D. J., Fernández, G., and Henson, R. N. (2012).How schema and novelty augment memory formation. Trends Neurosci. 35,211–219. doi: 10.1016/j.tins.2012.02.001Wang, J. Y., Weber, F. D., Zinke, K., Noack, H., and Born, J. (2017). Effects of sleepon word pair memory in children - separating item and sourcememory aspects.Front. Psychol. 8:1533. doi: 10.3389/fpsyg.2017.01533Wilhelm, I., Diekelmann, S., and Born, J. (2008). Sleep in children improvesmemory performance on declarative but not procedural tasks. Learn. Mem. 15,373–377. doi: 10.1101/lm.803708Conflict of Interest Statement: The authors declare that the research wasconducted in the absence of any commercial or financial relationships that couldbe construed as a potential conflict of interest.Copyright © 2018 Johnson and Durrant. This is an open-access article distributedunder the terms of the Creative Commons Attribution License (CC BY). The use,distribution or reproduction in other forums is permitted, provided the originalauthor(s) and the copyright owner are credited and that the original publicationin this journal is cited, in accordance with accepted academic practice. No use,distribution or reproduction is permitted which does not comply with these terms.Frontiers in Psychology | www.frontiersin.org 3 June 2018 | Volume 9 | Article 1022

Commentary: Effects of Sleep on Word Pair Memory in Children–Separating Item and Source Memory Aspects

University of Lincoln Institutional Repository

Author’s ProofCarefully read the entire proof and mark all corrections in the appropriate place, using the Adobe Reader commenting tools (Adobe Help). Donot forget to reply to the queries.We do not accept corrections in the form of edited manuscripts.In order to ensure the timely publication of your article, please submit the corrections within 48 hours.If you have any questions, please contact science.production.office@frontiersin.org.Author Queries FormQ1 The citation and surnames of all of the authors have been highlighted.Please check all of the names carefully and indicate if any areincorrect. Please note that this may affect the indexing of your articlein repositories such as PubMed.Q2 Confirm that the email address in your correspondence section isaccurate.Q3 Verify that all the equations and special characters are displayedcorrectly.Q4 Ensure, if it applies to your study, the ethics statement is included inthe article.Q5 Ensure to add all grant numbers and funding information, as afterpublication this is no longer possible.Q6 The author contribution section is mandatory and a standardstatement has been inserted. Please edit as needed to accuratelyreflect the contribution of each author.Q7 Please provide the name of the department for “University of Lincoln,Lincoln, United Kingdom.”Q8 Please add the following references to the reference list.“Giedd et al., 1999; Huttenlocher, 1990; Luna and Sweeney, 2004;Huber and Born, 2014.”Q9 Please reduce short running title to maximum of five words.123456789101112131415161718192021222324252627282930313233343536373839404142434445464748495051525354555657585960616263646566676869707172737475767778798081828384858687888990919293949596979899100101102103104105106107108109110111112113114GENERAL COMMENTARYpublished: xx June 2018doi: 10.3389/fpsyg.2018.01022Frontiers in Psychology | www.frontiersin.org 1 June 2018 | Volume 9 | Article 1022Edited by:Peter Klaver,University of Surrey, United KingdomReviewed by:Emma L. Axelsson,Uppsala University, SwedenJenny Richmond,University of New South Wales,Australia*Correspondence:Simon J. DurrantSiDurrant@lincoln.ac.ukQ2Specialty section:This article was submitted toDevelopmental Psychology,a section of the journalFrontiers in PsychologyReceived: 08 December 2017Accepted: 31 May 2018Published: xx June 2018Citation:Johnson JM and Durrant SJ (2018)Commentary: Effects of Sleep onWord Pair Memory inChildren–Separating Item and SourceMemory Aspects.Front. Psychol. 9:1022.doi: 10.3389/fpsyg.2018.01022Commentary: Effects of Sleep onWord Pair Memory inChildren–Separating Item and SourceMemory AspectsQ1Jennifer M. Johnson and Simon J. Durrant*University of Lincoln, Lincoln, United Kingdom Q7Keywords: declarative memory consolidation, sleep, children, hippocampus, prefrontal cortexA commentary onEffects of Sleep on Word Pair Memory in Children–Separating Item and Source MemoryAspectsby Wang, J. Y., Weber, F. D., Zinke, K., Noack, H., and Born, J. (2017). Front. Psychol. 8:1533.doi: 10.3389/fpsyg.2017.01533Evidence of sleep-dependent memory consolidation in adults has become widespread in the last Q4Q3two decades (Rasch and Born, 2013). However, investigations of similar consolidation in childrenare still at an early stage (Maski et al., 2015).Previous research has shown sleep-dependent memory consolidation in children on taskssuch as lexical integration of novel words (Henderson et al., 2012) and spatial location learning(Kurdziel et al., 2013). However, this appears to be limited to declarative memory tasks; procedurallearning tasks do not show the same sleep-related benefit in children that is seen in adults (Fischeret al., 2007; Wilhelm et al., 2008). Declarative memory tasks are strongly hippocampal-dependent(Squire, 1992), and consolidation of hippocampal memories is particularly associated with slowwave sleep (SWS) (Diekelmann et al., 2009; Born and Wilhelm, 2012). It has therefore beensuggested that the large amount of SWS seen during childhood (Ohayon et al., 2004) is the reasondeclarative memories show sleep-dependent consolidation in children.However, not all declarative memories are equal. In particular, a key distinction can be drawnbetween the memory of an object (item memory) and the context in which it was encountered(source memory). The recent paper byWang et al. (2017) used a clever design to separate these twoforms of memory, and examined their consolidation in children across an 11 h interval containingwake or sleep or a shorter 1 h wake interval, and compared this to adults with an 11 h sleep or1 h wake interval. The design used two lists of word pairs and the innovation was to measurememory not only for the target words using cued recall (item memory), but also for memory ofwhich list they came from (source memory). When comparing adults to children, a key finding wasthat memory for word pairs was lower after sleep in children (compared to before), but showed noreduction after sleep in adults (Figure 2B in the original paper). Interestingly, this turned out to bedriven entirely by the word pairs without source memory; where the item and source had both beenremembered, no difference was seen.The authors interpreted their findings in the context of less distinct encoding in childrenleading to more rapid unbinding of items and sources. However, this does not explain whyitem-only memory declines across sleep in children but item-source memory remains intact, andwhy the opposite pattern (stronger consolidation across sleep of item-only memory compared toitem-source memory) is seen in adults.115116117118119120121122123124125126127128129130131132133134135136137138139140141142143144145146147148149150151152153154155156157158159160161162163164165166167168169170171172173174175176177178179180181182183184185186187188189190191192193194195196197198199200201202203204205206207208209210211212213214215216217218219220221222223224225226227228Johnson and Durrant Commentary: Effects of Sleep on Word Pair Memory in Children–Separating Item and Source Memory AspectsWe believe that the influential model by Preston andEichenbaum (2013) allows a more comprehensive frameworkin which to explain the findings. This model postulates theexistence of a “what” stream of cortical processing fromthe dorsal hippocampus through the perirhinal cortex, anda “where” stream from the dorsal hippocampus through theparahippocampal cortex. Additionally, the ventral hippocampusconnects to the medial prefrontal cortex (mPFC), where pre-existing schemata influence subsequent memory processing (Tseet al., 2007; van Kesteren et al., 2010, 2012), and exert a top-down influence on “what” facilitating segregation from “where.”Importantly, Eichenbaum (2014, 2017) has suggested that the“where” stream also processes temporal context information ina similar way, i.e., it may also operate as a “when” stream.A key difference in the declarativememory systems of childrenand adults is that the density of hippocampal synapses in childrenreaches similar levels to adults within the first 6 months oflife (Seress and Ábrahám, 2008), but the frontal lobes havea prolonged developmental course (Luna and Sweeney, 2004)Q8with myelinization (Giedd et al., 1999), synaptic density countsQ8(Huttenlocher, 1990) and overall function of the prefrontalQ8cortex not reaching adult levels until late adolescence (Huberand Born, 2014). In children we should therefore expectQ8to see stronger bottom-up processing, and a reduced top-down influence of existing schemata, which would otherwisetypically strengthen item (“what”) memory at the expense of listcontext (“where/when”). This is supported by previous evidenceconfirming that more rapid consolidation of schematic memoriesoccurs during sleep and involves the mPFC (Durrant et al., 2015),and that sleep-dependent consolidation leads to a reduction inassociated contextual information (Cairney et al., 2011).For the Wang et al. (2017) study, greater decontextualizationand consolidation during sleep in adults compared to childrenwould show up as a bigger reduction in item-only memory inchildren and this is exactly what the authors found (Figure 2D).However, we would expect to see relatively little change acrossQ9sleep for items with their original contexts intact in both childrenand adults, reflecting equal hippocampal development, and wesee this too (Figure 2C). This argument is further strengthenedby the fact that the relationship between SWS and memoryconsolidation seen in the study in children was limited to itemswhich retained their context. It also explains why the childrenshowed little sleep-dependent benefit for items which have losttheir contextual cues (Figure 1F).In children, memory for the lists themselves was alsostronger after sleep compared to wake (Figure 1G), while inadults, no significant enhancement of list memory was seen(Supplementary Material). If memory for the lists makes use ofthe contextual stream, the absence of strong mPFC influenceallows both streams to consolidate during sleep, whereas thetop-down mPFC influence in adults strengthens the “what”stream at the expense of the “where/when” stream (hence listmemory is not enhanced after sleep compared to wake inadults).The neural basis of our interpretation could be tested in fMRI,where we predict that prefrontal activation will be stronger inadults during sleep and associated with item-only memory, whilehippocampal activation would be associated with both item andsource memory, but more strongly for items which retain theircontext.The Wang et al. (2017) paper has shown the clear benefitsof measuring different aspects of declarative memory in a singlestudy. We hope this new interpretation will help further improveour understanding of different memory systems and stimulateresearch in this area.Q5AUTHOR CONTRIBUTIONSAll authors listed have made a substantial, direct and intellectual Q6contribution to the work, and approved it for publication.REFERENCESBorn, J., and Wilhelm, I. (2012). System consolidation of memory during sleep.Psychol. Res. 76, 192–203. doi: 10.1007/s00426-011-0335-6Cairney, S. A., Durrant, S. J., Musgrove, H., and Lewis, P. A. (2011). Sleep andenvironmental context: interactive effects for memory. Exp. Brain Res. 214,83–92. doi: 10.1007/s00221-011-2808-7Diekelmann, S., Wilhelm, I., and Born, J. (2009). The whats and whensof sleep-dependent memory consolidation. Sleep Med. Rev. 13, 309–321.doi: 10.1016/j.smrv.2008.08.002Durrant, S. J., Cairney, S. A., McDermott, C., and Lewis, P. A. (2015). Schema-conformant memories are preferentially consolidated during REM sleep.Neurobiol. Learn. Mem. 122, 41–50. doi: 10.1016/j.nlm.2015.02.011Eichenbaum, H. (2014). Time cells in the hippocampus: A new dimension formapping memories. Nat. Rev. Neurosci. 15, 732–744. doi: 10.1038/nrn3827Eichenbaum, H. (2017). Time (and space) in the hippocampus. Curr. Opin. Behav.Sci. 17, 65–70. doi: 10.1016/j.cobeha.2017.06.010Fischer, S., Wilhelm, I., and Born, J. (2007). Developmental differences in sleep’srole for implicit off-line learning: comparing children with adults. J. Cogn.Neurosci. 19, 214–227. doi: 10.1162/jocn.2007.19.2.214Henderson, L. M., Weighall, A. R., Brown, H., and Gareth Gaskell, M. (2012).Consolidation of vocabulary is associated with sleep in children. Dev. Sci. 15,674–687. doi: 10.1111/j.1467-7687.2012.01172.xKurdziel, L., Duclos, K., and Spencer, R. M. (2013). Sleep spindles in middaynaps enhance learning in preschool children. Proc. Natl. Acad. Sci. U.S.A. 110,17267–17272. doi: 10.1073/pnas.1306418110Maski, K., Holbrook, H., Manoach, D., Hanson, E., Kapur, K., and Stickgold,R. (2015). Sleep dependent memory consolidation in children withautism spectrum disorder. Sleep 38, 1955–1963. doi: 10.5665/sleep.5248Ohayon, M. M., Carskadon, M. A., Guilleminault, C., and Vitiello, M. V. (2004).Meta-analysis of quantitative sleep parameters from childhood to old agein healthy individuals: developing normative sleep values across the humanlifespan. Sleep 27, 1255–1273. doi: 10.1093/sleep/27.7.1255Preston, A. R., and Eichenbaum, H. (2013). Interplay of hippocampusand prefrontal cortex in memory. Curr. Biol. 23, R764–R773.doi: 10.1016/j.cub.2013.05.041Rasch, B., and Born, J. (2013). About sleep’s role in memory. Physiol. Rev. 93,681–766. doi: 10.1152/physrev.00032.2012Seress, L., and Ábrahám, H. (2008). “Pre-and postnatal morphologicaldevelopment of the human hippocampal formation,” in Handbook ofDevelopmental Cognitive Neuroscience, 2nd Edn, eds C. A. Nelson and M.Luciana (Cambridge, MA: MIT Press), 187–209.Squire, L. R. (1992). Declarative and nondeclarative memory: multiple brainsystems supporting learning and memory. J. Cogn. Neurosci. 4, 232–243.doi: 10.1162/jocn.1992.4.3.232Frontiers in Psychology | www.frontiersin.org 2 June 2018 | Volume 9 | Article 1022229230231232233234235236237238239240241242243244245246247248249250251252253254255256257258259260261262263264265266267268269270271272273274275276277278279280281282283284285286287288289290291292293294295296297298299300301302303304305306307308309310311312313314315316317318319320321322323324325326327328329330331332333334335336337338339340341342Johnson and Durrant Commentary: Effects of Sleep on Word Pair Memory in Children–Separating Item and Source Memory AspectsTse, D., Langston, R. F., Kakeyama, M., Bethus, I., Spooner, P. A., Wood, E.R., et al. (2007). Schemas and memory consolidation. Science 316, 76–82.doi: 10.1126/science.1135935van Kesteren, M. T., Fernández, G., Norris, D. G., and Hermans, E.J. (2010). Persistent schema-dependent hippocampal-neocorticalconnectivity during memory encoding and postencoding rest in humans.Proc. Natl. Acad. Sci. U.S.A. 107, 7550–7555. doi: 10.1073/pnas.0914892107van Kesteren, M. T., Ruiter, D. J., Fernández, G., and Henson, R. N. (2012).How schema and novelty augment memory formation. Trends Neurosci. 35,211–219. doi: 10.1016/j.tins.2012.02.001Wang, J. Y., Weber, F. D., Zinke, K., Noack, H., and Born, J. (2017). Effects of sleepon word pair memory in children - separating item and sourcememory aspects.Front. Psychol. 8:1533. doi: 10.3389/fpsyg.2017.01533Wilhelm, I., Diekelmann, S., and Born, J. (2008). Sleep in children improvesmemory performance on declarative but not procedural tasks. Learn. Mem. 15,373–377. doi: 10.1101/lm.803708Conflict of Interest Statement: The authors declare that the research wasconducted in the absence of any commercial or financial relationships that couldbe construed as a potential conflict of interest.Copyright © 2018 Johnson and Durrant. This is an open-access article distributedunder the terms of the Creative Commons Attribution License (CC BY). The use,distribution or reproduction in other forums is permitted, provided the originalauthor(s) and the copyright owner are credited and that the original publicationin this journal is cited, in accordance with accepted academic practice. No use,distribution or reproduction is permitted which does not comply with these terms.Frontiers in Psychology | www.frontiersin.org 3 June 2018 | Volume 9 | Article 1022

Commentary: Effects of sleep on word pair memory in children–separating item and source memory aspects

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Commentary: Effects of sleep on word pair memory in children–separating item and source memory aspects

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