睡眠对婴幼儿学习的记忆巩固作用
The role of sleep in consolidating memory of learning in infants and toddlers
通讯作者: 张丹丹, E-mail:zhangdd05@gmail.com
收稿日期: 2023-06-15
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Received: 2023-06-15
睡眠依赖性记忆巩固指在睡眠期间, 大脑对新学习的信息或技能进行重新处理和加强, 从而使记忆更加稳定和持久的过程。睡眠在将新习得的信息巩固到稳定的长时记忆的过程中发挥了重要的作用。记忆类型不同, 睡眠依赖性记忆巩固的作用也有所不同。同时, 睡眠的不同阶段和特征对不同类型记忆巩固的影响也有差异。在成人研究的基础上, 近年的婴幼儿研究发现, 即使在个体发展的早期阶段, 睡眠也具有记忆巩固的重要作用。在学习后经历睡眠的婴幼儿与那些没有经历睡眠的控制组相比, 学习效果显著提高、可以更好更快地解决问题。婴幼儿在睡眠时, 海马、内侧颞叶等与记忆有关的脑区会显著激活, 睡眠纺锤波、慢波等脑电特征与婴幼儿记忆巩固效果相关。从陈述性记忆和程序性记忆两种不同的记忆类型入手, 介绍婴幼儿睡眠依赖性记忆巩固的行为和脑研究的进展, 帮助掌握睡眠对婴幼儿学习的记忆巩固作用。
关键词:
Sleep-dependent memory consolidation refers to the process during sleep in which the brain reprocesses and reinforces newly acquired information or skills, thereby enhancing the stability and longevity of memories. Sleep plays a pivotal role in consolidating recently acquired knowledge into enduring long-term memories. The influence of sleep-dependent memory consolidation varies depending on the type of memory, with different stages and characteristics of sleep exerting distinct effects on various memory processes.
Given the significant differences in sleep structure and physiological mechanisms between infants and adults, it is imperative not to extrapolate findings from adult studies directly to infants and toddlers. Additionally, owing to the remarkable neuroplasticity of the infant brain and its unique sleep patterns, investigating the impact of sleep on memory consolidation in infants can significantly deepen our comprehension of the neural mechanisms underlying sleep-dependent memory consolidation.
Building upon adult research, we present a synthesis of recent studies focusing on infants and toddlers, highlighting the critical role of sleep in memory consolidation during early development. Infants and toddlers who nap or sleep after learning consistently exhibit superior memory retention and enhanced problem-solving abilities compared to those who remain awake. During sleep, brain regions associated with memory, such as the hippocampus and medial temporal lobe, demonstrate significant activation. Distinct electroencephalogram (EEG) features, such as sleep spindles and slow waves, correlate with memory consolidation in infants and toddlers.
This paper addresses two primary forms of memory: declarative memory and procedural memory, shedding light on the impact of sleep-dependent memory consolidation in infants. In the realm of declarative memory, sleep enhances the quantity and accuracy of various episodic memory components, encompassing cartoon faces, toy manipulation, spatial locations, and chronological sequences. Moreover, distinct sleep features, such as sleep spindles and slow waves, make unique contributions to different episodic memories. Sleep also fosters selective memory consolidation, knowledge transfer, and the activation of memory-related brain regions, including the hippocampus, in infants and young children. These findings furnish valuable insights into the neural mechanisms governing sleep's role in early episodic memory development.
Regarding procedural memory, though limited studies exist on the relationship between infant sleep and procedural memory consolidation, some evidence suggests a positive influence of sleep on infant procedural memory. Future research should explore the interplay between sleep and motor skill development in infants, with particular emphasis on the role of rapid eye movement (REM) sleep, as adult studies underscore its significance in procedural memory consolidation.
Despite the progress in this field, several unresolved questions persist. Future research should aim to address whether sleep exerts a memory-consolidating effect on newborns, elucidate the distinctions between sleep-dependent memory consolidation in infants and adults, systematically investigate the impact of sleep on infants' social and language learning, and discern how different sleep types, durations, and timings in infants and young children contribute to memory consolidation.
Keywords:
本文引用格式
彭芝琳, 郑若颖, 胡晓晴, 张丹丹.
PENG Zhilin, ZHENG Ruoying, HU Xiaoqing, ZHANG Dandan.
1 引言
成人研究发现, 学习后的睡眠有助于巩固新学到的信息, 从而产生稳定和持久的记忆表征(Schmid et al., 2020; Stickgold, 2005)。记忆的形成包括学习以及随后的两个记忆巩固阶段——记忆稳定和记忆增强(也称为离线学习) (Walker, 2005)。最初的学习以及稳定(维持学习后记忆表征)阶段并不依赖于睡眠, 而增强(在无主动复述情况下提高记忆)阶段则通常发生在睡眠中(Al-Sharman & Siengsukon, 2014; Farhadian et al., 2021; Sugawara et al., 2014; Walker, 2005)。学习之后经历睡眠的个体其学习效果显著高于一直保持清醒的个体(Walker et al., 2002), 这是由学习后睡眠中出现的记忆重放现象对学习内容进行巩固造成的(Eichenlaub et al., 2020)。这种通过睡眠发生的记忆巩固效应称为睡眠依赖性记忆巩固(Stickgold, 2005)。
成人研究证明睡眠对陈述性和程序性记忆均具有巩固作用(Diekelmann et al., 2009; Schmid et al., 2020):与始终保持清醒相比, 学习后的睡眠不仅增强了陈述性记忆, 而且提高了程序性记忆的表现(Wang et al., 2021)。但对不同类型的记忆, 睡眠对记忆巩固的脑神经机制可能有所不同(Ackermann & Rasch, 2014)。研究发现, 陈述性记忆巩固会激活海马旁回以及丘脑、内侧颞叶、前额叶和小脑(van Dongen et al., 2012), 而程序性记忆巩固则与纹状体、海马、小脑和运动皮层的激活有关(Albouy et al., 2008; Cousins et al., 2016)。同时研究还发现, 陈述性记忆的巩固主要发生在非快速眼动(non-rapid eye movement, NREM)睡眠阶段(Rasch et al., 2007), 而程序性记忆的巩固则主要发生在快速眼动(rapid eye movement, REM)睡眠阶段(Plihal & Born, 1997; Qian et al., 2022)。同时成人记忆巩固研究表明, 睡眠纺锤波是重要的睡眠脑电指标之一, 睡眠纺锤波的出现频率和幅度或能量与大脑对新信息的巩固以及记忆保持密切相关(Ulrich, 2016)。
婴儿是指出生后1岁以内的孩子(包括新生儿期, 即出生后28天以内的婴儿); 幼儿是指1岁到3岁的孩子(有时也泛指1到6岁的孩子)。婴幼儿研究同样提示了睡眠对记忆巩固的促进作用。行为学研究表明, 睡眠能提高婴幼儿的学习能力(Berger & Scher, 2017; Seehagen et al., 2015), 后者主要指婴幼儿在发育过程获取新知识和技能的能力, 例如运动技能学习(爬行、坐立、站立和行走等)、感知觉学习(识别颜色和数量等) (Veldman et al., 2019)。神经影像学研究发现, 在婴幼儿最初学习中被激活的脑区会在随后的睡眠中被重新激活(Johnson et al., 2020)。脑电研究显示了婴幼儿特定类型的睡眠依赖性记忆巩固与睡眠结构特征之间的联系(Friedrich et al., 2019; Satomaa et al., 2020)。然而需要注意的是, 人类的睡眠在生命的最初几年经历了快速发展(Iglowstein et al., 2003; Jenni et al., 2007), 婴幼儿的睡眠模式与成人存在很大差异。在生命早期, 新生儿就表现出了多相睡眠模式, 包括3种睡眠类型:主动睡眠、安静睡眠和不确定睡眠(Ednick et al., 2009)。从出生后第2个月开始, 主动睡眠逐渐演变成REM睡眠, 安静睡眠演变成NREM睡眠(Jenni et al., 2004; Kurth et al., 2015)。成人的NREM睡眠和REM睡眠整晚交替进行, 大约90分钟一个周期(McCarley, 2007); 3月龄婴儿的这两种睡眠循环大约每60分钟发生一次(Davis et al., 2004)。在生命的第一年中, 睡眠纺锤波的密度、持续时间和频率等在额叶和顶叶均持续增加并达到整个生命的峰值(D'Atri et al., 2018; Jenni et al., 2004)。睡眠纺锤波在婴儿1~2月龄时出现(Grigg-Damberger et al., 2007), 而12~30月龄时睡眠纺锤波的功率、密度和持续时间开始下降(D'Atri et al., 2018; Page et al., 2018), 并在整个生命周期中持续下降(Clawson et al., 2016; Lokhandwala & Spencer, 2022)。睡眠慢波可在2~4月龄婴儿中观察到(Fattinger et al., 2014), 该脑电波在生命最初几年逐渐增加, 并在青春期前后开始减少(Campbell & Feinberg, 2009)。婴儿期丰富的睡眠纺锤波和睡眠慢波对早期记忆的编码和巩固都有重要作用。
此外与成人不同, 白天睡眠(即小睡)是婴幼儿期睡眠的常见组成部分(Galland et al., 2012; Iglowstein et al., 2003), 对生命早期记忆的发展至关重要(Horváth & Plunkett, 2018)。首先, 白天睡眠可以帮助婴幼儿巩固和提取白天学习的新信息, 增强记忆的稳定性和持久性。有研究发现, 6月龄和12月龄的婴儿在学习后4小时内进行30分钟以上的小睡可使记忆保存得更久(Seehagen et al., 2015)。其次, 白天睡眠可以强化婴幼儿的学习效果。研究显示, 婴幼儿在学习新技能后如果紧接着睡一觉, 其技能水平会显著提高(Berger & Scher, 2017)。这说明白天睡眠可以增强婴幼儿学习过程中形成的新记忆。
综上, 由于婴幼儿和成人在睡眠的结构和生理机制等诸多方面均有显著不同, 因此成人睡眠依赖性记忆巩固的发现不能直接推广到婴幼儿中。同时, 由于婴幼儿大脑的极强可塑性及其特殊的睡眠特征, 考察婴幼儿睡眠对记忆巩固的作用可进一步加深我们对睡眠依赖性记忆巩固脑机制的理解。本文就睡眠在婴幼儿记忆巩固过程中的作用进行综述, 分别从陈述性记忆和程序性记忆的睡眠巩固作用两个方面进行总结, 为深入了解睡眠对婴幼儿神经发育、认知发展、运动发展的影响提供理论基础。
2 睡眠对婴幼儿陈述性记忆的巩固
在整个婴幼儿期, 白天的小睡被认为在巩固陈述性记忆方面发挥着重要的作用(Mason et al., 2021)。陈述性记忆包括情景记忆(与时间或空间背景相关的自传式记忆)和语义记忆(对一般知识的记忆)。需要指出的是, 由于婴幼儿还没有完全发展出语义理解和概念化等语言能力, 因此在语义记忆部分, 我们将重点介绍睡眠对婴幼儿语言和概念泛化能力的影响。
2.1 睡眠对婴幼儿情景记忆的巩固
在婴幼儿睡眠的情景记忆研究中, Horváth等(2018)使用视觉配对比较任务考察了3月龄婴儿的睡眠依赖性记忆巩固, 发现只有学习后经过小睡(1.5小时)的婴儿才能记住学习过的卡通人脸, 并且婴儿学习的速度与睡眠脑电指标相关:婴儿对卡通人脸产生习惯化(habituation)所花的时间与额叶睡眠纺锤波的密度成反比。美国Spencer课题组使用两种不同的陈述性记忆任务考察了睡眠对3~6岁幼儿情景记忆的影响。第一项研究(Kurdziel et al., 2013)采用视觉空间学习任务, 发现经历小睡的幼儿对卡通图片位置的回忆准确率高于未经历小睡的幼儿, 且这种差异在24小时后仍然显著, 睡眠纺锤波的密度与回忆正确率成正比。第二项研究(Lokhandwala & Spencer, 2021)采用卡通图片序列记忆任务, 同样发现学习后睡眠组幼儿在延迟测试中的回忆正确率高于清醒组幼儿, 且组间差异在24小时后仍然显著, 回忆正确率与慢波睡眠时间呈正相关。Spencer的两项研究表明, 睡眠的不同特征对幼儿期的不同情景性记忆有独特的贡献:空间记忆可能更依赖于睡眠纺锤波, 而情景序列记忆可能更多依赖于睡眠慢波。
延迟模仿(deferred imitation)是一种被广泛认可的情景记忆的测量方法(McDonough et al., 1995), 该任务让被试对一段时间之前观看过的他人行为进行模仿。德国Seehagen课题组在3项研究中采用延迟模仿范式, 让婴儿面对面观看和学习记忆实验者手工操作玩具的过程(开小车、为玩偶更换手套等), 进而考察小睡对婴儿动作模仿记忆的作用。第一项研究(Seehagen et al., 2015)发现在动作模仿学习之后经过小睡, 6月龄和12月龄婴儿对所学动作的记忆数量和准确率均高于学习后未小睡的同年龄婴儿, 且这种组间差异能保持至少24小时。该研究提供了支持睡眠对一岁以内婴儿陈述性记忆巩固作用的首项实验证据。Seehagen课题组的第二项研究(Konrad, Herbert, et al., 2016)发现, 在动作模仿学习之后经过小睡, 12月龄婴儿不但对所学动作的记忆数量明显高于未经历小睡的婴儿, 而且能从呈现的新刺激中提取出之前的学习要点, 这表明睡眠能帮助婴儿将最近获得的知识应用到新环境中。Seehagen课题组的第三项研究(Konrad, Seehagen, et al., 2016)进一步将12月龄婴儿分为学习后睡眠组、清醒组和基线组(未经历学习), 且学习和测试阶段使用了颜色不同的玩具(例如将学习过的红色小鼠玩偶在测试阶段换成粉色的小鼠玩偶), 以考察学习和记忆的灵活性。结果发现睡眠组表现出对所学动作更高的记忆数量, 且比清醒组更快地执行首个学过的动作, 该实验表明睡眠可促进婴儿将已学知识灵活应用于类似线索的记忆检索。综上, Seehagen课题组的3项研究均表明睡眠对婴儿记忆编码和记忆巩固具有重要作用(Seehagen et al., 2019)。此外, Konrad等(2019)在另一项动作模仿研究中考察了幼儿的选择性睡眠记忆巩固。研究者在学习阶段按照一定顺序向15月龄和24月龄幼儿展示了手工操作4个玩具的相关动作(例如移除玩具车前的障碍让车驶下斜坡)和不相关动作(例如移除玩具车后的障碍, 车仍静止)共8个动作序列, 发现在学习后未经历睡眠的幼儿大脑仍保留了之前学习过的8个动作序列的信息, 而经历了睡眠的幼儿并没有表现出此种有序回忆的模式:他们只对4个玩具相关动作有明确的记忆。该研究首次证明了幼儿的选择性睡眠记忆巩固, 即睡眠可帮助幼儿有选择地“抛弃”在学习中他们认为对未来无用或不相关的信息。
目标记忆重激活(targeted memory reactivation, TMR)是考察睡眠依赖性记忆巩固神经机制的重要范式(Hu et al., 2020)。成人研究发现陈述性记忆的TMR过程显著激活海马旁回、丘脑和内侧颞叶等记忆相关脑区(van Dongen et al., 2012)。美国Ghetti课题组采用TMR范式分别考察了2岁幼儿的睡眠对情景记忆和语义记忆巩固的神经机制。在第一项情景记忆研究中(Prabhakar et al., 2018), 实验者让幼儿在特定房间与特定玩偶玩耍并收听目标歌曲, 然后在学习后一周的自然夜间睡眠时, 播放目标歌曲和新异歌曲并采集功能磁共振信号。结果发现, 当在睡眠中播放与情景记忆相关的记忆线索时(目标歌曲, 正常播放和逆时序播放两种条件), 与播放新异歌曲相比, 幼儿双侧海马的激活增加, 且右侧海马的激活与幼儿对目标歌曲与房间位置和玩偶的关联记忆准确度呈正相关。该研究提示了海马在早期发育过程中对情景记忆巩固的重要作用。Ghetti课题组的第二项情景记忆研究(Mooney et al., 2021)让幼儿按照先后顺序学习3个人物与3个地点的配对(首先医生去了学校, 接着消防员去了滑梯, 最后宇航员去了校园巴士), 一旦幼儿能按时间顺序成功配对出3对人物−空间关系, 实验程序就为幼儿播放一首奖励性歌曲(目标歌曲)。结果发现幼儿在学习后延迟一周的测试中仍能较好地记住空间位置信息, 但对时间顺序的记忆成绩较差。学习一周后在睡眠期间播放目标歌曲, 发现目标歌曲(与新异歌曲相比)激活的右侧海马神经活动水平与幼儿的时间顺序记忆准确度呈正相关, 而与空间位置记忆成绩无显著相关, 这揭示了海马与时间顺序记忆之间的紧密联系。上述两项研究的海马激活结果在空间记忆条件不一致, 这可能是研究所采用的记忆任务不同引起的。第一项研究使用了单一的空间位置配对任务, 第二项研究由于同时考察空间位置记忆和时间顺序记忆, 使用了更复杂的序列记忆任务。两项研究的学习任务有差异, 可能导致记忆巩固的神经机制有所不同。我们建议未来能有更多的TMR实验进一步考察海马与时间顺序记忆以及空间位置记忆之间的关系。
总之, 上述研究发现, 睡眠能提高婴幼儿对卡通人脸、玩具操作、空间位置和时间顺序等不同类型的情景记忆的记忆数量和准确率, 且睡眠的不同特征(如纺锤波和慢波)对不同情景记忆有独特的贡献。此外, 睡眠还能促进婴幼儿的选择性记忆巩固和知识迁移, 同时激活海马等与记忆相关脑区。这些研究为探索睡眠对早期发育过程中情景记忆的神经机制提供了有价值的信息。
2.2 睡眠对婴幼儿语义记忆及泛化能力的影响
与情景记忆类似, 白天的小睡对婴幼儿语义记忆巩固的作用也非常重要。在一项观察性研究中(Horváth & Plunkett, 2016), 研究者通过让家长填写睡眠日记来收集并记录8~36月龄婴幼儿的白天和夜间睡眠模式。结果发现婴幼儿白天小睡的次数以及夜间睡眠效率与4个月后幼儿产出性词汇(指学习者在口语或书面语表达中能自主使用的词汇)和接受性词汇(指学习者能理解其最基本词义的词汇)的增长均呈正相关, 而夜间睡眠的长度不提供词汇增长的预测信息。
在婴幼儿语言学习的实验室研究中, 美国的Nadel课题组在两项实验中采用习惯化−转头偏好范式, 考察了睡眠对15月龄幼儿语言学习的影响。研究者让幼儿先学习按特定拼读规则生成的24个人工“伪词”的语音, 然后在幼儿小睡后测试他们对所学单词的记忆。第一项研究(Gómez et al., 2006)发现, 学习之后的小睡有助于幼儿从已学习的单词范例中抽提出拼读规则, 并将此规则灵活应用于没有学习过的单词。第二项研究(Hupbach et al., 2009)发现, 幼儿只有在学习后进行了小睡, 才能在学习语音范例24小时后仍保持对拼读规则的记忆。以上两项研究结果表明睡眠对幼儿语言学习和记忆具有重要影响。此外, Simon等(2017)也采用习惯化−转头偏好范式考察了睡眠对6.5月龄婴儿单词概率学习的影响, 发现只有学习后经过小睡的婴儿才能记住语言的统计规律, 并且记忆巩固的效果与脑电节律的alpha和theta能量以及额−中央区的慢波能量均呈正相关。除了习惯化范式, Williams和Horst (2014)采用故事阅读任务考察了睡眠对3岁幼儿单词学习的影响。研究者让一组幼儿阅读同一个故事3遍而另一组幼儿阅读3个不同的故事, 目的是学习故事中出现的新单词, 之后再对两个阅读组进行二次分组, 让一半幼儿进行小睡, 另一半幼儿保持清醒。最后在学习后的2.5小时、24小时和一周后进行3次记忆测试。结果在两个阅读组别中均发现睡眠组幼儿对新单词的记忆准确度显著高于清醒组, 说明睡眠对单词学习具有记忆巩固作用; 此外重复阅读相同故事的幼儿比阅读不同故事的幼儿对单词的学习和记忆效果更好。
物体−类别任务是考察婴幼儿语义学习和泛化能力的最常用范式。德国的Friedrich课题组利用该范式并结合脑电指标(N400和纺锤波), 在系列研究中考察了睡眠对婴儿从语言感知到语义学习阶段的记忆重组和泛化作用。第一项研究(Friedrich et al., 2015)发现, 9~16月龄婴幼儿的小睡(约1.5小时)可以重组记忆并产生新的语义知识, 并且这种语义泛化效应(给新物体贴上学习过的类别标签)与额、顶、中央区睡眠纺锤波的幅度呈正相关。第二项研究(Friedrich et al., 2017)发现, 6~8月龄婴儿在学习了物体−类别配对之后, 只有经历了NREM睡眠才能产生语义泛化效应, 且该效应与NREM阶段的时长和睡眠纺锤波的幅度均呈正相关; 而浅睡(未经历NREM睡眠阶段)婴儿只能巩固语音感知相关的记忆。随后该课题组采用物体−类别范式并结合脑电的N400指标考察了14~17月龄幼儿睡眠中语义记忆和情景记忆的相互关系(Friedrich et al., 2020)。N400是大脑中央−顶区的负向波形, 由语义期待的违背所诱发。Friedrich等(2020)首先让幼儿学习物体与正确类别标签配对(先后呈现物体和类别标签), 这个过程形成了情景语境。接着让一组幼儿小睡而另一组保持清醒, 最后在测试阶段向幼儿呈现学习过的物体(旧物体)和新物体, 每个物体分别呈现两次, 一次与正确类别标签配对, 另一次与错误类别标签配对。结果发现, 新物体−错误类别标签在所有幼儿大脑诱发了N400(与正确配对条件相比), 表明幼儿通过语义学习形成了语义记忆。旧物体−错误类别标签仅在清醒组(而不是睡眠组)幼儿大脑诱发了N400(与正确配对条件相比); 而旧物体−正确标签配对在睡眠组幼儿的额−颞区诱发了比旧物体−错误标签配对幅度更大负波(反映了情景记忆效应), 且该脑电成分的幅度与额叶睡眠纺锤波的幅度呈正比。上述结果表明, 幼儿同时进行情景和语义学习后, 睡眠可以巩固情景记忆并保护精确的情景记忆暂时性地免受语义记忆的干扰。此外, 还有研究(Spanò et al., 2018)采用物体−类别范式考察REM睡眠对典型发育和唐氏综合症幼儿单词语义学习的影响。结果发现典型发育的1~4岁幼儿只有经历过小睡才能在记忆后测中表现出较高的准确率, 且睡眠的记忆巩固效应能至少保持24小时; 同时REM阶段的时长与单词学习准确率呈正相关。相反, 唐氏综合症患儿在经历小睡后单词语义学习的准确率与学习后保持清醒组相比有所下降, 且学习效果与REM阶段的时长无关。这是目前少有的将REM睡眠与陈述性记忆巩固联系起来的证据, 其结论的可靠性还需要后续研究的检验。
物体命名学习任务(又称为快速映射范式)与物体−类别任务相似, 指被试将特定物体与一个新词(名称)相联系的过程。Horst在英国和澳大利亚的课题组采用物体命名学习任务考察小睡对2.5岁幼儿语义学习的影响。第一项研究(Axelsson et al., 2018)通过向幼儿呈现新物体图片和新单词音频, 让幼儿将新物体和新单词进行配对来学习单词。在测试阶段让幼儿根据学习过的物体名称音频在屏幕上指出对应的物体图片。结果发现, 学习后睡眠组幼儿在学习后4小时和24小时的记忆测试中均表现出较高的、稳定的回忆准确率, 而清醒组幼儿在两次测试中的回忆准确率均显著低于睡眠组幼儿, 且学习后24小时的测试成绩比学习后4小时测试时明显降低。该研究表明小睡有助于维持并巩固幼儿对物体命名学习的记忆。该课题组第二项研究(Axelsson et al., 2021)在学习阶段增加了记忆强化环节, 即每轮物体−名称快速映射学习重复进行两次。研究结果与前一项研究相似, 且发现增加了记忆强化环节的幼儿在两次延迟测试(学习后4小时和24小时)中的成绩均高于学习后即时测试的成绩, 该发现突出了小睡对物体命名学习的记忆增强作用。
同样是物体命名学习任务, 但在测试时不让幼儿根据名称的语音在屏幕上指出对应的物体, 而是在听到名称时观测他们对目标物体和分心物体的注视时间, 这个指标被称为跨模态注视偏好(intermodal preferential looking, IPL; Golinkoff et al., 1987)。英国的Horváth课题组采用IPL指标在两项研究中考察了16月龄幼儿对新单词的学习情况。第一项研究(Horváth et al., 2015)在学习阶段让幼儿记住两个物体的语音标签。学习结束后将幼儿随机分为睡眠组和清醒组, 并在学习后即刻和2小时延迟后进行两次测试。结果发现在学习后即刻测试时两组幼儿的表现无差异, 而在睡眠组经历小睡后的第二次测试中, 睡眠组幼儿在听到名称标签后对目标物体的注视时间比清醒组幼儿更长。该课题组的第二项研究(Horváth et al., 2016)考察了小睡对幼儿词义泛化能力的影响。幼儿在学习阶段熟悉两个物体及其语音标签, 随后进行即刻的记忆测试。之后将幼儿随机分为睡眠组和清醒组, 在1.5小时后再次测试记忆。在两次测试中, 实验者向幼儿呈现两个旧物体的泛化形式(即与学习时颜色不同但形状相同的物体)以及未学习过的新物体。结果在IPL指标上发现了睡眠组别和测试次数的交互作用:睡眠组在小睡后听到标签时对目标物体的注视时间显著增加(与学习后即刻测试相比), 而清醒组对目标物体的注视时间在两次测试中差异不显著。该研究表明了小睡对幼儿语义泛化能力的作用, 即小睡是一个提取和保留语义概念关键特征的主动的语言表征增强过程。
前文提及的目标记忆重激活(TMR)范式也可用于考察语义记忆巩固的神经机制。除了情景记忆, Ghetti课题组也采用TMR范式考察了2岁幼儿的大脑是如何在睡眠中巩固新学到的单词(Johnson et al., 2021)。研究先让幼儿通过物体−标签任务学习单词(即物体命名的学习), 并在学习后即刻以及一周后对学过的单词进行记忆测试。结果显示, 幼儿学会了新单词, 并在一周后仍能保持对新学单词的记忆。在一周后的自然夜间睡眠时播放记忆线索(学习过的单词)时, 幼儿左侧海马和左侧前内侧颞叶被激活(与未学过的新单词相比), 且这些脑区的激活水平与幼儿对学过单词的记忆准确度和3岁的词汇产出量均呈正相关。该研究证明了以海马为核心的内侧颞叶在语义记忆巩固中的作用。
表1 睡眠对婴幼儿陈述性记忆的巩固
| 论文信息 | 设计类型及样本量 | 被试 年龄 | 实验任务 | 记忆测试时间 | 睡眠类型 | 主要发现 |
|---|---|---|---|---|---|---|
| Axelsson et al., First Language, 2018 | 混合设计, 40 | 2.5岁 | 快速映射 | 即时、4小时延迟、24小时延迟 | 白天小睡、夜间睡眠 | 回忆正确率:睡眠组>清醒组 |
| Axelsson et al., Brain Sciences, 2021 | 混合设计, 40 | 2.5岁 | 快速映射 | 即时、5小时延迟、24小时延迟 | 白天小睡、夜间睡眠 | 回忆正确率:睡眠组>清醒组 |
| Friedrich et al., Nature Communications, 2015 | 混合设计, 90 | 16、19月 | 物体−类别 | 1.5小时延迟 | 白天小睡 | 睡眠组出现N400、N200-500 |
| Friedrich et al., Current Biology, 2017 | 混合设计, 107 | 6~8月 | 物体−类别 | 1小时延迟 | 白天小睡 | 浅睡组出现晚期正成分、深睡组出现N400 |
| Friedrich et al., Nature Communications, 2020 | 混合设计, 60 | 14~17月 | 物体−类别 | 1小时延迟 | 白天小睡 | 睡眠组N400消失, 出现FTMR |
| Gómez et al., Psychological Science, 2006 | 混合设计, 48 | 15月 | 习惯化−转头偏好 | 4小时延迟 | 白天小睡 | 注视时间:睡眠组<控制组<清醒组 |
| Hupbach et al., Developmental Science, 2009 | 组内设计, 两个实验各24 | 15月 | 习惯化−转头偏好 | 24小时延迟 | 实验1:白天小睡, 实验2:夜间睡眠 | 注视时间:睡眠组<清醒组 |
| Horváth et al., Journal of Sleep Research, 2015 | 混合设计, 31 | 16月 | 跨模态注视偏好 | 即时、2小时延迟 | 白天小睡 | 注视时间:睡眠组<清醒组 |
| Horváth et al., Sleep, 2016 | 混合设计, 28 | 16月 | 跨模态注视偏好 | 即时、1.5小时延迟 | 白天小睡 | 注视时间:睡眠组<清醒组 |
| Horváth et al., Developmental Science, 2018 | 组间设计, 45 | 3月 | 视觉配对比较 | 1.5小时延迟 | 白天小睡 | 注视时间:睡眠组<清醒组 |
| Johnson et al., Current Biology, 2021 | 组内设计, 28 | 2岁 | 目标记忆重激活 | 即时、一周后延迟 | 夜间睡眠 | 左侧海马、左侧前内侧颞叶激活:目标单词>新异单词 |
| Kurdziel et al., PANS, 2013 | 混合设计, 40 | 3~6岁 | 视觉空间学习 | 即时、5小时延迟、24小时延迟 | 白天小睡、夜间睡眠 | 回忆正确率:睡眠组>清醒组 |
| Konrad et al., Neurobiology of Learning and Memory, 2016 | 混合设计, 60 | 12月 | 延迟模仿 | 4小时延迟、24小时延迟 | 白天小睡、夜间睡眠 | 回忆正确率:睡眠组>清醒组>基线组 |
| Konrad et al., Developmental Psychobiology, 2016 | 混合设计, 48 | 12月 | 延迟模仿 | 4小时延迟 | 白天小睡、夜间睡眠 | 回忆正确率:睡眠组>清醒组>基线组 |
| Konrad et al., Journal of Sleep Research, 2019 | 混合设计, 96 | 15、24月 | 延迟模仿 | 24小时延迟 | 白天小睡、夜间睡眠 | 无关动作回忆正确率:基线组<睡眠组<清醒组 |
| Lokhandwala & Spencer, Developmental Science, 2021 | 混合设计, 22 | 3~6岁 | 卡通图片序列记忆 | 即时、4小时延迟、24小时延迟 | 白天小睡、夜间睡眠 | 回忆正确率:睡眠组>清醒组 |
| Mooney et al., Developmental Cognitive Neuroscience, 2021 | 组内设计, 48 | 2岁 | 目标记忆重激活 | 一周后延迟 | 夜间睡眠 | 右侧海马激活:目标歌曲>新异歌曲 |
| Prabhakar et al., PNAS, 2018 | 组内设计, 22 | 2岁 | 目标记忆重激活 | 一周后延迟 | 夜间睡眠 | 双侧海马激活:目标歌曲>新异歌曲 |
| Seehagen et al., PNAS, 2015 | 组间设计, 实验1和2为120和96 | 6、12月 | 延迟模仿 | 实验1:4小时延迟; 实验2:24小时延迟 | 实验1:白天小睡, 实验2:夜间睡眠 | 回忆正确率:睡眠组>清醒组>基线组 |
| Simon et al., Brain Language, 2017 | 混合设计, 37 | 6.5月 | 习惯化−转头偏好 | 1小时延迟 | 白天小睡 | 注视时间:睡眠组<控制组<清醒组 |
| Spanò et al. PNAS, 2018 | 混合设计, 50 | 1~4岁 | 物体−类别 | 1小时延迟 | 白天小睡 | 睡眠组N400消失, 出现FTMR |
| Williams & Horst, Frontiers in Psychology, 2014 | 混合设计, 48 | 3岁 | 故事阅读 | 即时、2.5小时延迟、24小时延迟、一周后延迟 | 白天小睡、夜间睡眠 | 回忆正确率:睡眠组>清醒组 |
3 睡眠对婴幼儿程序性记忆的巩固
程序性记忆也称为技能记忆, 指我们对技巧、习惯和行为的记忆, 这些技能来自于学习和经验, 不一定能被有意识地回忆起来。研究程序性学习在婴幼儿期是具有挑战性的, 因为婴幼儿的运动能力尚处于发展阶段。目前关于婴幼儿睡眠与程序性记忆巩固的研究较少, 下文分别介绍。
最早的一项研究(Fagen & Rovee-Collier, 1983)训练3月龄婴儿通过踢腿的动作来移动婴儿床上方的手机。两周后再次展示手机, 发现踢腿率(即踢腿动作的保留概率)与婴儿在这两周期间的睡眠时间呈正相关。Gibson等(2011)考察了睡眠对12月龄婴儿运动发展的影响, 通过活动记录仪、睡眠日记和简明婴幼儿睡眠问卷来记录婴儿连续一周的睡眠状况, 并通过年龄和发育阶段问卷(ages and stages questionnaire, ASQ)来衡量婴儿的发展情况。结果发现, 婴儿夜间睡眠效率与ASQ问卷中解决问题技能和精细运动能力的得分均呈正相关, 表明睡眠可促进婴儿的技巧和运动发展。DeMasi等(2023)将10~19月龄的婴幼儿分为高低行走经验两组, 并通过佩戴活动记录仪来测量他们睡眠期间的身体活动。结果发现, 高行走经验组婴儿在睡眠期间有更多不规律的运动, 且身体活动次数逐小时递增。该研究表明婴儿夜间睡眠期间身体活动随着运动技能的学习而发生变化, 行走经验对婴儿睡眠和睡眠期间的运动有显著的影响。因此, 婴幼儿睡眠与运动是相互影响、相互促进的, 这对处于发展早期个体的认知发展有重要作用。在脑观测指标上, 婴幼儿程序性记忆的研究还很少, 但有初步证据表明睡眠纺锤波和慢波睡眠活动与精细运动能力的学习和巩固有关。例如Satomaa等(2020)发现在8月龄婴儿的夜间睡眠中, 左侧额叶和枕叶的睡眠慢波与精细运动能力呈正相关; Page等(2018)发现12~30月龄婴幼儿的精细运动能力与NREM睡眠中theta节律活动呈正相关, 与delta节律呈负相关。
隧道任务为考察婴幼儿的运动问题解决能力提供了一个很好的范式(Brawn et al., 2008)。在此任务中, 婴儿被放置在一条隧道前, 他们需要改变身体姿势(从站立变成爬行)才能进入隧道。婴儿在训练阶段通过非言语提示(在隧道另一端通过玩具吸引婴儿注意)学会爬行通过隧道。婴幼儿的运动表现可通过在任务中的姿势切换次数、通过隧道的时间以及需要提示的数量来测量。美国的Berger课题组在3项研究中采用该任务考察了睡眠对婴儿运动学习的影响。第一项研究(Berger & Scher, 2017)让9~16月龄的婴幼儿学习隧道任务, 比较学习后经历和未经历小睡的婴幼儿对运动问题的解决, 发现睡眠组比清醒组需要更少的提示就能通过隧道。第二项研究(DeMasi et al., 2021)将10~19月龄的婴幼儿分为两组, 其中一组在学习后立即小睡, 另一组在学习后延迟4小时再进行小睡, 发现在学习后6小时的测试中, 立即小睡组比延迟小睡组需要更少的提示和时间就能通过隧道, 且进入隧道前的姿势切换次数更少, 这凸显了学习后及时睡眠对婴幼儿程序性记忆巩固的重要作用。第三项研究(Horger et al., 2023)将10~19月龄的婴幼儿随机分为3组:学习和首次测试之间小睡组, 学习和首次测试之间保持清醒组和学习后立即测试组, 3组被试都在经历自然夜间睡眠后再次测试。结果发现, 学习和首次测试之间小睡组和清醒组在第一次测试中的隧道前姿势切换次数差异不显著, 但第二次测试时睡眠组的姿势切换次数显著低于清醒组, 表明只有动作学习后立即小睡的婴幼儿, 才能进一步通过自然夜间睡眠巩固程序性记忆从而提高他们在运动任务中的表现。此外, 本研究分别考察了白天小睡和夜间睡眠对婴幼儿程序性记忆的作用, 为后续研究提供了很好的借鉴思路。
序列反应时任务(serial reaction time task, SRT)是研究运动学习能力的经典范式, 该任务要求被试学习按照一定的顺序迅速按下若干按钮。Wilhelm等(2012)采用SRT考察睡眠对4~6岁幼儿程序性记忆的巩固作用。首先将幼儿分为高低强度学习两组(分别接受10次和30次训练), 所有幼儿在学习后经历两次测试(30分钟延迟测试、2小时延迟测试), 并在两次测试之间进行小睡或保持清醒。结果发现只有经过高强度学习后, 小睡组幼儿在2小时延迟测试中的反应时才显著低于清醒组幼儿; 而睡眠对SRT反应时的影响在低强度学习条件下不显著。该研究表明睡眠对程序性记忆的巩固作用还受到睡前幼儿程序性学习水平的影响。Desrochers等(2016)采用SRT考察不同类型睡眠对3~6岁幼儿程序性记忆的影响, 所有幼儿在学习后经历3次测试(学习后即刻、5小时延迟测试、24小时延迟测试)。结果发现在5小时延迟测试中, 小睡组幼儿的反应时和正确率改善与学习后即刻测试时没有显著差异, 但在24小时后(经历了夜间睡眠), 基于反应时和正确率测量的行为能力显著提高。与Horger等(2023)的发现一致, 本研究也提示程序性学习必须要通过夜间睡眠才能形成稳定的记忆表征。此外, 有研究采用SRT及其变式以考察睡眠障碍对8~11岁儿童的程序性记忆巩固的影响(Csábi et al., 2016), 结果发现与健康儿童相似, 睡眠呼吸暂停综合征患儿在程序性学习后经过自然夜间睡眠, 显著改善了序列按键的反应时和正确率, 这表明睡眠对患有睡眠障碍儿童的程序性记忆巩固也具有积极作用。
表2 睡眠对婴幼儿程序性记忆的巩固
| 论文信息 | 设计类型 及样本量 | 被试年龄 | 实验任务 | 记忆测试时间 | 睡眠类型 | 主要发现 |
|---|---|---|---|---|---|---|
| Berger & Scher, Journal of Experimental Child Psychology, 2017 | 混合设计, 28 | 9~16月 | 隧道任务 | 即时、6小时延迟 | 白天小睡 | 提示数量:睡眠组<清醒组 |
| Csábi et al., Frontiers in Human Neuroscience, 2016 | 混合设计, 32 | 8~11岁 | 序列反应时 | 即时、12小时延迟 | 夜间睡眠 | 回忆正确率:睡眠组>清醒组; 反应时:睡眠组<清醒组 |
| DeMasi et al., Infant Behavior & Development, 2021 | 混合设计, 29 | 10~19月 | 隧道任务 | 6小时延迟 | 白天小睡 | 提示数量:立即小睡组<延迟小睡组; 姿势切换数量:立即小睡组<延迟小睡组 |
| DeMasi et al., Infancy, 2023 | 被试间设计, 78 | 10~19月 | 无 | 无 | 夜间睡眠 | 不规律的运动数量:高行走经验组>低行走经验组 |
| Desrochers et al., Experimental Brain Research, 2016 | 混合设计, 36 | 3~6岁 | 序列反应时 | 即时、5小时延迟、24小时延迟 | 白天小睡、夜间睡眠 | 反应时:睡眠组<清醒组 |
| Fagen & Rovee-Collier, Science, 1983 | 被试内设计, 16 | 3月 | 踢腿任务 | 连续两周 | 夜间睡眠 | 踢腿率与睡眠时间呈正相关 |
| Gibson et al., Sleep and Biological Rhythms, 2011 | 被试内设计, 52 | 12月 | 无 | 连续一周 | 夜间睡眠 | 夜间睡眠效率与解决问题技能和精细运动能力呈正相关 |
| Wilhelm et al., Developmental Science, 2012 | 混合设计, 35 | 4~6岁 | 序列反应时 | 30分钟延迟、2小时延迟 | 白天小睡 | 反应时:睡眠组<清醒组 |
| Satomaa et al., Sleep, 2020 | 混合设计, 36 | 8月 | 无 | 无 | 夜间睡眠 | 左侧额叶和枕叶的睡眠慢波与精细运动能力呈正相关 |
| Page et al., Sleep, 2018 | 混合设计, 36 | 12~30月 | 无 | 无 | 夜间睡眠 | 精细运动能力与theta节律呈正相关, 与delta节律呈负相关 |
4 总结与展望
通过对已有文献的回顾, 我们发现睡眠对婴幼儿陈述性和程序性记忆均有重要的巩固作用, 其中白天小睡能显著帮助陈述性记忆的巩固; 而相比于陈述性记忆, 程序性记忆巩固对夜间睡眠的依赖程度似乎更高。婴幼儿在睡眠时, 与记忆有关的脑区(如海马、内侧颞叶等)会被激活, 同时纺锤波、慢波振幅等睡眠脑电特征的变化也与婴幼儿记忆巩固效果相关。尽管如此, 目前该领域仍有一些问题未解决, 我们建议未来围绕以下几个问题开展研究:
第一, 睡眠对新生儿是否也有记忆巩固作用?目前关于睡眠记忆巩固的研究多集中在成人和儿童青少年(Mason et al., 2021; Schmid et al., 2020), 至今报道的睡眠依赖性记忆巩固发生的最小年龄为3月龄(Fagen & Rovee-Collier, 1983; Horváth et al., 2018), 还未有研究考察过睡眠对新生儿学习的记忆巩固作用。已知年龄越小, 大脑的可塑性越强。新生儿刚刚脱离母体来到全新的世界, 他们每天都在接收周围环境中的信息, 并且通过学习建立大量突触连接, 形成陈述性和程序性记忆表征。睡眠在这一过程中是否以及起到了什么作用?新生儿虽然表现出了成人睡眠周期的雏形, 但尚未发展出NREM和REM等不同睡眠阶段。我们建议未来利用脑电、近红外等技术实时监测新生儿睡眠状态, 并结合TMR范式考察睡眠和相应的神经活动对新生儿学习的记忆巩固作用。
第二, 婴幼儿和成人的睡眠依赖性记忆巩固有何异同?睡眠可以促进大脑产生新的突触以形成稳定的记忆表征(Yang et al., 2014)。突触数量的变化是大脑对经验进行学习和适应的过程, 也是记忆巩固的重要机制之一。人类个体从出生开始, 大脑神经元突触的数量会经历先增加后减少的发展变化趋势, 具体的, 1岁以前是突触数量发展的关键时期, 3岁时突触数量达到顶峰(大约是成人的两倍), 之后逐渐开始修剪, 青春期16岁左右突触数量达到稳定。婴幼儿和成人在突触数量以及发育阶段的不同, 可能会导致二者睡眠依赖性记忆巩固的机制也有所不同, 因此不能简单地将成人的研究结论推广至婴幼儿。目前在成人的研究中对于睡眠与记忆巩固关系的解释, 主要包括突触稳态理论、系统巩固假说(Squire & Alvarez, 1995)和记忆片段重演重叠理论(Lewis & Durrant, 2011)。研究婴幼儿的睡眠与记忆巩固关系, 不仅可以帮助我们了解婴幼儿的认知发展和学习能力, 也可以为成人的理论提供新的视角和证据。例如, 婴幼儿睡眠是否也存在海马和新皮层之间的信息交流?婴幼儿的突触是否也会在睡眠中进行优化和删减?婴幼儿的记忆片段是否也会在睡眠中重演?这些问题都值得进一步探索。考虑到目前成人研究的大部分实验范式都无法在婴幼儿被试中使用, 未来研究需要探索新的适合婴幼儿的实验范式, 从而在新范式中对比婴幼儿和成人的睡眠依赖性记忆巩固机制。例如, 我们建议选用对婴幼儿简单且便于他们行为输出的学习和记忆测试任务, 包括但不限于视觉辨别学习任务(可考察眼动指标), 双耳竞争性语音学习任务(可考察转头偏好等), 或者采用oddball范式结合脑电MMR指标等, 直接利用脑神经指标评估婴幼儿被试的记忆表现。
第三, 进一步系统考察睡眠对婴幼儿社会和语言学习有何促进作用?与儿童青少年和成人要进行大量的陈述性和程序性学习(例如知识, 技能)不同, 婴幼儿阶段最主要的任务是进行社会学习(包括安全依恋、人际互动、情绪、心理理论)和语言学习(母语, 语音语调, 词汇等) (Bremner & Fogel, 2004; Slot et al., 2020; Tomasello, 2001)。尽管已有研究探讨了睡眠对婴幼儿语音、语义、卡通人脸等学习记忆的影响, 但还有许多社会和语言学习的方面没有涉及。例如, 不少研究强调动物和人类早期经验的重要性, 生物学家甚至提出了印刻的观点(Robledo et al., 2022)。那么婴幼儿睡眠对于社会学习的形成有何促进作用?哪些学习经验会优先被睡眠巩固加工, 从而形成知识图式, 进而影响之后的认知和社会发展?又如, 有成人研究发现睡眠能降低负性情绪的强度, 甚至能增强正性情绪体验, 以及提高对他人面孔、情绪的记忆(Tempesta et al., 2018; Walker & van der Helm, 2009)。那么, 睡眠对婴幼儿的情绪和社会记忆(社会性微笑、陌生人焦虑、分离焦虑等)是否也有特别的作用?我们建议未来研究采用适合于婴幼儿的社会和语言学习的实验范式, 考察睡眠对婴幼儿社会和语言记忆(例如对母亲和陌生人面孔、声调和语调的记忆)的巩固作用。
第四, 婴幼儿睡眠的类型、时长和睡眠的时间点等是否对记忆巩固有不同的贡献?目前的研究表明, 白天小睡对婴幼儿的陈述性记忆有促进作用, 程序性记忆巩固对夜间睡眠的依赖程度较高, 然而仍缺乏对睡眠类型进行直接比较的研究。白天小睡和夜间睡眠是否对不同记忆类型有特异性的影响?进一步的, 白天小睡和夜间睡眠对记忆的不同影响是否源于NREM/REM在两种睡眠中的占比不同?此外, 睡眠的时长(例如2小时和5小时的小睡相比)以及睡眠开始的时间(例如学习后即刻入睡和学习后2小时再入睡相比)对记忆巩固的影响如何?这些都是值得探讨的问题。
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There is increasing evidence that sleep promotes off-line enhancement of a variety of explicitly learned motor tasks in young adults. However, whether sleep promotes off-line consolidation of implicitly learned motor tasks is still under question. Furthermore, the role of sleep in promoting transfer of learning remains unknown. This study examined the role of sleep in learning and transfer of learning of an implicit continuous motor task. Twenty-three neurologically intact individuals (mean age 26.4 years) were randomly assigned to either a sleep group or a no-sleep group. The sleep group practiced a continuous tracking task in the evening and underwent retention and transfer testing the following morning, while the no-sleep group practiced the tracking task in the morning and underwent retention and transfer testing in the evening. The results show that in both the sleep and no-sleep groups, performance improved off-line without further practice for both the general skill and the sequence-specific skill. The results also indicate that sleep and time promote transfer of learning of both sequence-specific and general skill learning to a spatial and temporal variation of the motor task. These findings demonstrate that sleep does not play a critical role in promoting off-line learning and transfer of learning of an implicit continuous motor task.
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It is now recognized that extensive maturational changes take place in the human brain during adolescence, and that the trajectories of these changes are best studied longitudinally. We report the first longitudinal study of the adolescent decline in non-rapid eye movement (NREM) delta (1-4 Hz) and theta (4-8 Hz) EEG. Delta and theta are the homeostatic frequencies of human sleep. We recorded sleep EEG in 9- and 12-year-old cohorts twice yearly over a 5-year period. Delta power density (PD) was unchanged between age 9 and 11 years and then fell precipitously, decreasing by 66% between age 11 and 16.5 years (P <.000001). The decline in theta PD began significantly earlier than that in delta PD and also was very steep (by 60%) between age 11 and 16.5 years (P <.000001). These data suggest that age 11-16.5 years is a critically important maturational period for the brain processes that underlie homeostatic NREM EEG, a finding not suggested in previous cross-sectional data. We hypothesize that these EEG changes reflect synaptic pruning. Comparing our data with published longitudinal declines in MRI-estimated cortical thickness suggests the theta age curve parallels the earlier maturational thinning in 3-layer cortex, whereas the delta curve tracks the later changes in 5-layer cortex. This comparison also reveals that adolescent declines in NREM delta and theta are substantially larger than decreases in cortical thickness (>60% vs. <20%). The magnitude, interindividual difference, and tight link to age of these EEG changes indicate that they provide excellent noninvasive tools for investigating neurobehavioral correlates of adolescent brain maturation.
Form and function of sleep spindles across the lifespan
Cued reactivation of motor learning during sleep leads to overnight changes in functional brain activity and connectivity
Declarative and non-declarative memory consolidation in children with sleep disorder
Different maturational changes of fast and slow sleep spindles in the first four years of life
DOI:S1389-9457(17)31583-6
PMID:29458750
[本文引用: 2]
Massive changes in brain morphology and function in the first years of life reveal a postero-anterior trajectory of cortical maturation accompanied by regional modifications of NREM sleep. One of the most sensible marker of this maturation process is represented by electroencephalographic (EEG) activity within the frequency range of sleep spindles. However, direct evidence that these changes actually reflect maturational modifications of fast and slow spindles still lacks. Our study aimed at answering the following questions: 1. Do cortical changes at 11.50 Hz frequency correspond to slow spindles? 2. Do fast and slow spindles show different age trajectories and different topographical distributions? 3. Do changes in peak frequency explain age changes of slow and fast spindles?We measured the antero-posterior changes of slow and fast spindles in the first 60 min of nightly sleep of 39 infants and children (0-48 mo.).We found that (A) changes of slow spindles from birth to childhood mostly affect frontal areas (B) variations of fast and slow spindles across age groups go in opposite direction, the latter progressively increasing across ages; (C) this process is not merely reducible to changes of spindle frequency.As a main finding, our cross-sectional study shows that the first form of mature spindle (i.e., corresponding to the adult phasic event of NREM sleep) is marked by the emergence of slow spindles on anterior regions around the age of 12 months.Copyright © 2017 Elsevier B.V. All rights reserved.
Sleep in infants and young children: Part one: Normal sleep
The importance of sleep to overall health and well-being is becoming increasingly appreciated; however, clinicians may not have a sound understanding of the fundamentals of sleep. This review of normal pediatric sleep is meant to provide a foundation for the pediatric nurse practitioner to develop and use in clinical practice. Key concepts such as normal sleep physiology including biological rhythms and stages of sleep are discussed. Developmental changes in sleep seen in the transition from infancy to young childhood are highlighted, and strategies for instituting and maintaining normal sleep behaviors are recommended. Part 2 of this series will address common sleep problems experienced by young children.
Nap timing makes a difference: Sleeping sooner rather than later after learning improves infants' locomotor problem solving
Infant motor development predicts the dynamics of movement during sleep
DOI:10.1111/infa.v28.2 URL [本文引用: 2]
Delayed benefit of naps on motor learning in preschool children
DOI:10.1007/s00221-015-4506-3
PMID:26645305
[本文引用: 2]
Sleep benefits memory consolidation across a variety of domains in young adults. However, while declarative memories benefit from sleep in young children, such improvements are not consistently seen for procedural skill learning. Here we examined whether performance improvements on a procedural task, although not immediately observed, are evident after a longer delay when augmented by overnight sleep (24 h after learning). We trained 47 children, aged 33-71 months, on a serial reaction time task and, using a within-subject design, evaluated performance at three time points: immediately after learning, after a daytime nap (nap condition) or equivalent wake opportunity (wake condition), and 24 h after learning. Consistent with previous studies, performance improvements following the nap did not differ from performance improvements following an equivalent interval spent awake. However, significant benefits of the nap were found when performance was assessed 24 h after learning. This research demonstrates that motor skill learning is benefited by sleep, but that this benefit is only evident after an extended period of time.
The whats and whens of sleep-dependent memory consolidation
DOI:10.1016/j.smrv.2008.08.002
PMID:19251443
[本文引用: 1]
Sleep benefits memory consolidation. The reviewed studies indicate that this consolidating effect is not revealed under all circumstances but is linked to specific psychological conditions. Specifically, we discuss to what extent memory consolidation during sleep depends on the type of learning materials, type of learning and retrieval test, different features of sleep and the subject population. Post-learning sleep enhances consolidation of declarative, procedural and emotional memories. The enhancement is greater for weakly than strongly encoded associations and more consistent for explicitly than implicitly encoded memories. Memories associated with expected reward gain preferentially access to sleep-dependent consolidation. For declarative memories, sleep benefits are more consistently revealed with recall than recognition procedures at retrieval testing. Slow wave sleep (SWS) particularly enhances declarative memories whereas rapid eye movement (REM) sleep preferentially supports procedural and emotional memory aspects. Declarative memory profits already from rather short sleep periods (1-2 h). Procedural memory profits seem more dose-dependent on the amount of sleep following the day after learning. Children's sleep with high amounts of SWS distinctly enhances declarative memories whereas elderly and psychiatric patients with disturbed sleep show impaired sleep-associated consolidation often of declarative memories. Based on the constellation of psychological conditions identified we hypothesize that access to sleep-dependent consolidation requires memories to be encoded under control of prefrontal-hippocampal circuitry, with the same circuitry controlling subsequent consolidation during sleep.
A review of the effects of sleep during the first year of life on cognitive, psychomotor, and temperament development
DOI:10.1093/sleep/32.11.1449
PMID:19928384
[本文引用: 1]
During the first year of life, infants spend most of their time in the sleeping state. Assessment of sleep during infancy presents an opportunity to study the impact of sleep on the maturation of the central nervous system (CNS), overall functioning, and future cognitive, psychomotor, and temperament development. To assess what is currently known regarding sleep during infancy and its effects on cognitive, psychomotor, and temperament development, we assessed the relevant literature published over the last several decades. To provide a foundation for a more in-depth understanding of this literature, we preface this with an overview of brain maturation, sleep development, and various assessment tools of both sleep and development during this unique period. At present, we do not have sufficient data to conclude that a causal relationship exists between infant sleep and cognitive, psychomotor, and temperament development. Caution should be used in predicting outcomes, as the timing and subjectivity of evaluations may obviate accurate assessment. Collectively, studies assess a wide array of sleep measures, and findings from one developmental period cannot be generalized readily to other developmental periods. Future studies should follow patients longitudinally. Additionally, refinements of existing assessment tools would be useful. In view of the relatively high reported pediatric prevalence of cognitive and behavioral deficits that carry significant long-term costs to individuals and society, early screening of sleep-related issues may be a useful tool to guide targeted prevention and early intervention.
Replay of learned neural firing sequences during rest in human motor cortex
Understanding sleep problems in children with epilepsy: Associations with quality of life, Attention-Deficit Hyperactivity Disorder and maternal emotional symptoms
DOI:10.1016/j.seizure.2016.06.011
PMID:27394056
[本文引用: 1]
This study aimed to (1) compare sleep problems between children and adolescents with epilepsy and non-epileptic controls, and (2) examine whether there is an association between sleep problems and quality of life, Attention-Deficit Hyperactivity Disorder (ADHD) and mothers' emotional symptoms.Fifty-three patients from a cohort of epilepsy (aged 7-18 years) and 28 controls with minor medical problems (aged 7-18 years) were included. Parents completed Children's Sleep Habits Questionnaire (CSHQ) and Kinder Lebensqualitätsfragebogen: Children's Quality of Life Questionnaire-revised (KINDL-R) for patients and controls. Turgay DSM-IV Disruptive Behavior Disorders Rating Scale (T-DSM-IV-S) parent and teacher forms were used to assess ADHD symptoms for patients. Mothers of the patients completed Beck Depression Inventory and State-Trait Anxiety Inventory (STAI). Neurology clinic charts were reviewed for the epilepsy-related variables.Children with epilepsy had a higher CSHQ Total score than the control group. Those with a CSHQ score >56 (which indicates moderate to severe sleep problems) had lower scores on KINDL-R. Parent-rated T-DSM-IV-S Total and Hyperactivity-Impulsivity scores, STAI trait and Beck scores were found to be higher in those with a CSHQ score >56. Significant positive correlations were found between CSHQ Total score and T-DSM-IV-S, STAI trait and Beck scores. Binary logistic regression analysis revealed that T-DSM-IV-S Total, Inattention and Hyperactivity-Impulsivity scores were significantly associated with a higher CSHQ Total score. None of the epilepsy-related variables were found to be related with the CSHQ Total score.Among children with epilepsy, sleep problems lead to a poor quality of life. The link between sleep problems and psychiatric symptoms must be conceptualized as a bilateral relationship. ADHD appears to be the strongest predictor of sleep problems.Copyright © 2016 British Epilepsy Association. Published by Elsevier Ltd. All rights reserved.
Memory retrieval: A time-locked process in infancy
Three-month-old infants learned to activate an overhead crib mobile by operant footkicking and received a visual reminder of the event (a "reactivation treatment") 2 weeks later, after forgetting had occurred. Subsequent manifestation of the association was a monotonic increasing function of time since the reactivation treatment, and performance of infants tested 8 hours after the remainder was related to the time spent sleeping in the interim (r = 0.75). These data demonstrate that normal retrieval is time-dependent. Moreover, individual data suggest that retrieval may be continuous rather than discontinuous.
The role of daytime napping in declarative memory performance: A systematic review
DOI:10.1016/j.sleep.2021.05.019
PMID:34148000
[本文引用: 1]
Sleep plays an important role in stabilizing and reinforcing memory of newly acquired information. Like nocturnal sleep, a daytime nap is shown to effectively contribute to memory processing. However, studies are often focused on nocturnal sleep. This review has aimed at systematically compiling the results of studies which have examined the effects of napping on declarative memory performance in healthy adults. Such studies have focused on different aspects of memory reinforcement following a diurnal nap including the involved mechanisms in memory reconsolidation, type of declarative tasks, cross-gender differences, the role of age, duration of nap and its delayed onset. One of the reviewed studies reported that even as short as 6 min of napping exerts a positive effect on memory function. Evidence from these studies indicates hippocampal-dependent enhancement of the learned information. Diurnal naps predominantly include non-rapid eye movement sleep with slow waves yielding potential effects on declarative memory. Evidence has shown that the empowered learning and retrieval depends upon spindle density during the nap. Moreover, the role of coordinated autonomic and central events in enhancing declarative memory has also been reported. Slow waves and sleep spindles are known to fuel declarative memory function during the NREM-2 (N2) stage of sleep.Copyright © 2021 Elsevier B.V. All rights reserved.
Overnight changes in the slope of sleep slow waves during infancy
DOI:10.5665/sleep.3390
PMID:24497653
[本文引用: 1]
Slow wave activity (SWA, 0.5-4.5 Hz) is a well-established marker for sleep pressure in adults. Recent studies have shown that increasing sleep pressure is reflected by an increased synchronized firing pattern of cortical neurons, which can be measured by the slope of sleep slow waves. Thus we aimed at investigating whether the slope of sleep slow waves might provide an alternative marker to study the homeostatic regulation of sleep during early human development.All-night sleep electroencephalography (EEG) was recorded longitudinally at 2, 4, 6, and 9 months after birth.Home recording.11 healthy full-term infants (5 male, 6 female).None.The slope of sleep slow waves increased with age. At all ages the slope decreased from the first to the last hour of non rapid-eye-movement (NREM) sleep, even when controlling for amplitude differences (P < 0.002). The decrease of the slope was also present in the cycle-by-cycle time course across the night (P < 0.001) at the age of 6 months when the alternating pattern of low-delta activity (0.75-1.75 Hz) is most prominent. Moreover, we found distinct topographical differences exhibiting the steepest slope over the occipital cortex.The results suggest an age-dependent increase in synchronization of cortical activity during infancy, which might be due to increasing synaptogenesis. Previous studies have shown that during early postnatal development synaptogenesis is most pronounced over the occipital cortex, which could explain why the steepest slope was found in the occipital derivation. Our results provide evidence that the homeostatic regulation of sleep develops early in human infants.
The reciprocal relation between sleep and memory in infancy: Memory-dependent adjustment of sleep spindles and spindle-dependent improvement of memories
Sleep-dependent memory consolidation in infants protects new episodic memories from existing semantic memories
Generalization of word meanings during infant sleep
DOI:10.1038/ncomms7004
PMID:25633407
[本文引用: 2]
Friedrich, Manuela Humboldt Univ, Dept Psychol, D-12489 Berlin, Germany. Friedrich, Manuela; Friederici, Angela D. Max Planck Inst Human Cognit & Brain Sci, Dept Neuropsychol, D-04103 Leipzig, Germany. Wilhelm, Ines Univ Childrens Hosp, Child Dev Ctr, CH-8032 Zurich, Switzerland. Wilhelm, Ines; Born, Jan Univ Tubingen, Inst Med Psychol & Behav Neurobiol, D-72076 Tubingen, Germany. Wilhelm, Ines; Born, Jan Univ Tubingen, Ctr Integrat Neurosci, D-72076 Tubingen, Germany.
The sleeping infant brain anticipates development
DOI:S0960-9822(17)30807-2
PMID:28756948
[本文引用: 2]
From the age of 3 months, infants learn relations between objects and co-occurring words [1]. These very first representations of object-word pairings in infant memory are considered as non-symbolic proto-words comprising specific visual-auditory associations that can already be formed in the first months of life [2-5]. Genuine words that refer to semantic long-term memory have not been evidenced prior to 9 months of age [6-9]. Sleep is known to facilitate the reorganization of memories [9-14], but its impact on the perceptual-to-semantic trend in early development is unknown. Here we explored the formation of word meanings in 6- to 8-month-old infants and its reorganization during the course of sleep. Infants were exposed to new words as labels for new object categories. In the memory test about an hour later, generalization to novel category exemplars was tested. In infants who took a short nap during the retention period, a brain response of 3-month-olds [1] was observed, indicating generalizations based on early developing perceptual-associative memory. In those infants who napped longer, a semantic priming effect [15, 16] usually found later in development [17-19] revealed the formation of genuine words. The perceptual-to-semantic shift in memory was related to the duration of sleep stage 2 and to locally increased sleep spindle activity. The finding that, after the massed presentation of several labeled category exemplars, sleep enabled even 6-month-olds to create semantic long-term memory clearly challenges the notion that immature brain structures are responsible for the typically slower lexical development.Copyright © 2017 Elsevier Ltd. All rights reserved.
Normal sleep patterns in infants and children: A systematic review of observational studies
DOI:10.1016/j.smrv.2011.06.001
PMID:21784676
[本文引用: 1]
This is a systematic review of the scientific literature with regard to normal sleep patterns in infants and children (0-12 years). The review was conducted according to the Preferred Reporting Items for Systematic Reviews and Meta-Analyses (PRISMA) guidelines. Mean and variability data for sleep duration, number of night wakings, sleep latency, longest sleep period overnight, and number of daytime naps were extracted from questionnaire or diary data from 34 eligible studies. Meta-analysis was conducted within age-bands and categories. In addition, fractional polynomial regression models were used to estimate best-fit equations for the sleep variables in relation to age. Reference values (means) and ranges (±1.96 SD) for sleep duration (hours) were: infant, 12.8 (9.7-15.9); toddler/preschool, 11.9 (9.9-13.8); and child, 9.2 (7.6-10.8). The best-fit (R(2)=0.89) equation for hours over the 0-12 year age range was 10.49-5.56×[(age/10)^0.5-0.71]. Meta-regression showed predominantly Asian countries had significantly shorter sleep (1h less over the 0-12 year range) compared to studies from Caucasian/non-Asian countries. Night waking data provided 4 age-bands up to 2 years ranging from 0 to 3.4 wakes per night for infants (0-2 months), to 0-2.5 per night (1-2 year-olds). Sleep latency data were sparse but estimated to be stable across 0-6 years. Because the main data analysis combined data from different countries and cultures, the reference values should be considered as global norms.Copyright © 2011 Elsevier Ltd. All rights reserved.
Actigraphic sleep and developmental progress of one-year-old infants
DOI:10.1111/sbr.2012.10.issue-2 URL [本文引用: 2]
The eyes have it: Lexical and syntactic comprehension in a new paradigm
Naps promote abstraction in language-learning infants
DOI:10.1111/j.1467-9280.2006.01764.x
PMID:16913948
[本文引用: 2]
Infants engage in an extraordinary amount of learning during their waking hours even though much of their day is consumed by sleep. What role does sleep play in infant learning? Fifteen-month-olds were familiarized with an artificial language 4 hr prior to a lab visit. Learning the language involved relating initial and final words in auditory strings by remembering the exact word dependencies or by remembering an abstract relation between initial and final words. One group napped during the interval between familiarization and test. Another group did not nap. Infants who napped appeared to remember a more abstract relation, one they could apply to stimuli that were similar but not identical to those from familiarization. Infants who did not nap showed a memory effect. Naps appear to promote a qualitative change in memory, one involving greater flexibility in learning.
The visual scoring of sleep and arousal in infants and children
Age is probably the single most crucial factor determining how humans sleep. Age and level of vigilance significantly influence the electroencephalogram (EEG) and the polysomnogram (PSG). The Pediatric Task Force provide an evidence-based review of the age-related development of the polysomnographic features of sleep in neonates, infants, and children, assessing the reliability and validity of these features, and assessing alternative methods of measurement. We used this annotated supporting text to develop rules for scoring sleep and arousals in infants and children. A pediatric EEG or PSG can only be determined to be normal by assessing whether the EEG patterns are appropriate for maturational age. Sleep in infants at term can be scored as NREM and REM sleep because all the polysomnographic and EEG features of REM sleep are present and quiet sleep, if not NREM sleep, is at least "not REM sleep." The dominant posterior rhythm (DPR) of relaxed wakefulness increases in frequency with age: (1) 3.5-4.5 Hz in 75% of normal infants by 3-4 months post-term; (2) 5-6 Hz in most infants 5-6 months post-term; 3) 6 Hz in 70% of normal children by 2 months of age; and 3) 8 Hz (range 7.5-9.5 Hz) in 82% of normal children age 3 years, 9 Hz in 65% of 9-year-olds, and 10 Hz in 65% of 15-year-old controls. Sleep spindles in children occur independently at two different frequencies and two different scalp locations: 11.0-12.75 Hz over the frontal and 13.0-14.75 Hz over the centroparietal electrodes; these findings are most prominent in children younger than 13 years. Centroparietal spikes are often maximal over the vertex (Cz), less often maximal over the left central (C3) or right central (C4) EEG derivation. About 50% of sleep spindles within a particular infant's PSG are asynchronous before 6 months of age, 30% at 1 year. Based on this, we recommend that: (1) sleep spindles be scored as a polysomnographic signature of NREM stage 2 sleep (N2) at whatever age they are first seen in a PSG, typically present by 2 to 3 months post-term; (2) identify and score sleep spindles from the frontal and centroparietal EEG derivations, especially in infants and children younger than 13 years. NREM sleep in an infant or child can be scored if the dominant posterior rhythm occupies <50% of a 30-second epoch, and one or more of the following EEG patterns appear: (1) a diffuse lower voltage mixed frequency activity; (2) hypnagogic hypersynchrony; (3) rhythmic anterior theta of drowsiness; (4) diffuse high voltage occipital delta slowing; (5) runs or bursts of diffuse, frontal, frontocentral, or occipital maximal rhythmic 3-5 Hz slowing; (6) vertex sharp waves; and/or (7) post-arousal hypersynchrony. K complexes first appear 5 months post-term and are usually present by 6 months post-term, whereas clearly recognizable vertex sharp waves are most often seen 16 months post-term. Vertex sharp waves are best seen over the central (Cz, C3, C4) and K complexes over the frontal (Fz, F3, F4) electrodes. Slow wave activity (SWA) of slow wave sleep (SWS) is first seen as early as 2 to 3 months post-term and is usually present 4 to 4.5 months post-term. SWA of SWS in an infant or child often has a peak-to-peak amplitude of 100 to 400 microV. Based on consensus voting we recommended scoring N1, N2, and N3 corresponding to NREM 1, 2, and SWS whenever it was recognizable in an infant's PSG, usually by 4 to 4.5 months post-term (as early as 2-3 months post-term). Epochs of NREM sleep which contain no sleep spindles, K complexes, or SWA would be scored as N1; those which contain either K complexes or sleep spindles and <20% SWS as N2, and those in which >20% of the 30-second epoch contain 0.5 to 2 Hz >75 microV (usually 100-400 microV) activity as N3. The DPR should be scored in the EEG channel that is best observed, (typically occipital), but DPR reactive to eye opening can be seen in central electrodes. Because sleep spindles occur independently over the frontal and central regions in children, they should be scored whether they occur in the frontal or central regions. Because sleep spindles are asynchronous before age 2 years, simultaneous recording of left and right frontal and central activity may be warranted in children 1-2 years of age. Simultaneous recording of left, right, and midline central electrodes may be appropriate because of the asynchronous nature of sleep spindles before age 2 years, but reliability testing is needed. Evidence has shown that the PSG cannot reliably be used to identify neurological deficits or to predict behavior or outcome in infants because of significant diversity of results, even in normal infants. Normal sleep EEG patterns and architecture are present in the first year of life, even in infants with severe neurological compromise. Increasing evidence suggests that sleep and its disorders play critical roles in the development of healthy children and healthy adults thereafter. Reliability studies comparing head-to-head different scoring criteria, recording techniques, and derivations are needed so that future scoring recommendations can be based on evidence rather than consensus opinion. We need research comparing clinical outcomes with PSG measures to better inform clinicians and families exactly what meaning a PSG has in evaluating a child's suspected sleep disorder.
In pre-school children, sleep objectively assessed via sleep- EEGs remains stable over 12 months and is related to psychological functioning, but not to cortisol secretion
DOI:10.1016/j.jpsychires.2013.08.007
PMID:24011863
[本文引用: 1]
Cross-sectional studies provide evidence that in pre-schoolers poor sleep is by this age already associated with both poor psychological functioning and an increased cortisol secretion. However, long-term studies on the stability of sleep in pre-schoolers are scarce. The aim of the present study was therefore to investigate objectively assessed sleep in pre-schoolers longitudinally, and to predict sleep, psychological functioning and cortisol secretion prospectively as a function of sleep 12 months earlier.A total of 58 pre-schoolers (mean age: 5.43 years; 47% females) were re-assessed 12 months later (mean age: 6.4 years). Sleep-EEG recordings were performed, saliva cortisol was analysed, and parents and experts rated children's psychological functioning.Longitudinally, poor objective sleep at age 5.4 years was associated with poor objective sleep and psychological difficulties but not cortisol secretion 12 months later. At age 6.4 years, poor sleep was concurrently associated with greater psychological difficulties and increased cortisol secretion.In pre-schoolers, poor sleep objectively assessed at the age of 5.4 years was associated with poor sleep and psychological difficulties one year later. Data indicate that in pre-schoolers, sleep remains stable over a 12-months-period. Pre-schoolers with poor sleep appear to be at risk for developing further psychological difficulties.Copyright © 2013 Elsevier Ltd. All rights reserved.
In pre-school children, sleep objectively assessed via actigraphy remains stable over 12 months and is related to psychological functioning, but not to cortisol secretion
DOI:10.1016/j.jpsychires.2014.04.008
PMID:24814637
[本文引用: 1]
Studies of the long-term stability of sleep in pre-schoolers are scarce. The aim of the present study was to investigate objectively assessed sleep via actigraphy in pre-schoolers longitudinally, and to predict sleep, psychological functioning and cortisol secretion prospectively as a function of sleep 12 months earlier.A total of 73 pre-schoolers (mean age: 5.45 years; 53% females) were assessed again after 12 (mean age: 6.4 years). Sleep-actigraphy recordings were performed, saliva cortisol was analysed, and parents and experts rated children's psychological functioning.Longitudinally, poor sleep at age 5.45 years was associated with poor sleep and internalizing and peer problems but not with externalizing problems and hyperactivity, and cortisol secretion 12 months later. At age 6.4 years and cross-sectionally, poor sleep was concurrently associated with greater psychological difficulties and increased cortisol secretion.In pre-schoolers, poor sleep objectively assessed at age five was associated with psychological difficulties and poor sleep as assessed via actigraph and one year later. Results indicate that in pre-schoolers sleep remains stable over a 12-mont interval. Pre-schoolers with poor sleep appear to be at risk for developing further psychological difficulties.Copyright © 2014. Published by Elsevier Ltd.
The unique contributions of day and night sleep to infant motor problem solving
DOI:10.1016/j.jecp.2022.105536 URL [本文引用: 2]
Memory in 3-month-old infants benefits from a short nap.
A daytime nap facilitates generalization of word meanings in young toddlers
DOI:10.5665/sleep.5348
PMID:26237777
[本文引用: 2]
One of the key processes in language development is generalization--the selection and extension of relevant features and information to similar objects and concepts. Little is known about how sleep influences generalization, and studies on the topic are inconclusive. Our aim was to investigate how a nap affects generalization in 16-mo-olds. We hypothesized that a nap is necessary for successful generalization of word meanings.Twenty-eight 16-mo-old, typically developing toddlers were randomly assigned to nap and wake groups. We trained toddlers with two novel object-word pairs and tested their initial ability to generalize. Toddlers took part in an intermodal preferential looking task, in which they were shown different colored versions of the original objects and heard one of the trained labels. If toddlers understand the label, they are expected to increase their looking time to the target. Looking behavior was measured with an automated eye tracker. Afterward, the nap group went to sleep, while the wake group stayed awake for approximately 2 h. We then repeated the test of their performance on the generalization task.A significant interaction of group and session was found in preferential looking. The performance of the nap group increased after the nap, whereas that of the wake group did not change.Our results suggest that napping improves generalization in toddlers.© 2016 Associated Professional Sleep Societies, LLC.
Napping facilitates word learning in early lexical development
DOI:10.1111/jsr.12306
PMID:25950233
[本文引用: 2]
Little is known about the role that night-time sleep and daytime naps play in early cognitive development. Our aim was to investigate how napping affects word learning in 16-month-olds. Thirty-four typically developing infants were assigned randomly to nap and wake groups. After teaching two novel object-word pairs to infants, we tested their initial performance with an intermodal preferential looking task in which infants are expected to increase their target looking time compared to a distracter after hearing its auditory label. A second test session followed after approximately a 2-h delay. The delay contained sleep for the nap group or no sleep for the wake group. Looking behaviour was measured with an automatic eye-tracker. Vocabulary size was assessed using the Oxford Communicative Development Inventory. A significant interaction between group and session was found in preferential looking towards the target picture. The performance of the nap group increased after the nap, whereas that of the wake group did not change. The gain in performance correlated positively with the expressive vocabulary size in the nap group. These results indicate that daytime napping helps consolidate word learning in infancy. © 2015 European Sleep Research Society.
Frequent daytime naps predict vocabulary growth in early childhood
Spotlight on daytime napping during early childhood
DOI:10.2147/NSS.S126252
PMID:29576733
[本文引用: 2]
Daytime napping undergoes a remarkable change in early childhood, and research regarding its relationship to cognitive development has recently accelerated. In this review, we summarize our current understanding of this relationship focusing on children aged <5 years. First, we evaluate different studies on the basis of the experimental design used and the specific cognitive processes they investigate. Second, we analyze how the napping status of children may modulate the relationship between learning and napping. Third, the possible role of sleep spindles, ie, specific electroencephalographic components during sleep, in cognitive development is explored. We conclude that daytime napping is crucial in early memory development.
Promoting memory consolidation during sleep: A meta- analysis of targeted memory reactivation
DOI:10.1037/bul0000223 URL [本文引用: 1]
Nap-dependent learning in infants
DOI:10.1111/j.1467-7687.2009.00837.x
PMID:19840054
[本文引用: 2]
Sleep has been shown to aid a variety of learning and memory processes in adults (Stickgold, 2005). Recently, we showed that infants' learning also benefits from subsequent sleep such that infants who nap are able to abstract the general grammatical pattern of a briefly presented artificial language (Gomez, Bootzin & Nadel, 2006). In the present study, we demonstrate, for the first time, long-term effects of sleep on memory for an artificial language. Fifteen-month-old infants who had napped within 4 hours of language exposure remembered the general grammatical pattern of the language 24 hours later. In contrast, infants who had not napped shortly after being familiarized with the language showed no evidence of remembering anything about the language. Our findings support the view that infants' frequent napping plays an essential role in establishing long-term memory.
Sleep duration from infancy to adolescence: Reference values and generational trends
DOI:10.1542/peds.111.2.302
PMID:12563055
[本文引用: 2]
The main purpose of the present study was to calculate percentile curves for total sleep duration per 24 hours, for nighttime and for daytime sleep duration from early infancy to late adolescence to illustrate the developmental course and age-specific variability of these variables among subjects.A total of 493 subjects from the Zurich Longitudinal Studies were followed using structured sleep-related questionnaires at 1, 3, 6, 9, 12, 18, and 24 months after birth and then at annual intervals until 16 years of age. Gaussian percentiles for ages 3 months to 16 years were calculated for total sleep duration (time in bed) and nighttime and daytime sleep duration. The mean sleep duration for ages 1 to 16 years was estimated by generalized additive models based on the loess smoother; a cohort effect also had to be included. The standard deviation (SD) was estimated from the loess smoothed absolute residuals from the mean curve. For ages 3, 6, and 9 months, an alternative approach with a simple model linear in age was used. For age 1 month, empirical percentiles were calculated.Total sleep duration decreased from an average of 14.2 hours (SD: 1.9 hours) at 6 months of age to an average of 8.1 hours (SD: 0.8 hours) at 16 years of age. The variance showed the same declining trend: the interquartile range at 6 months after birth was 2.5 hours, whereas at 16 years of age, it was only 1.0 hours. Total sleep duration decreased across the studied cohorts (1974-1993) because of increasingly later bedtime but unchanged wake time across decades. Consolidation of nocturnal sleep occurred during the first 12 months after birth with a decreasing trend of daytime sleep. This resulted in a small increase of nighttime sleep duration by 1 year of age (mean 11.0 +/- 1.1 hours at 1 month to 11.7 +/- 1.0 hours at 1 year of age). The most prominent decline in napping habits occurred between 1.5 years of age (96.4% of all children) and 4 years of age (35.4%).Percentile curves provide valuable information on developmental course and age-specific variability of sleep duration for the health care professional who deals with sleep problems in pediatric practice.
Development of the nocturnal sleep electroencephalogram in human infants
Sleep duration from ages 1 to 10 years: Variability and stability in comparison with growth
DOI:10.1542/peds.2006-3300
URL
[本文引用: 1]
OBJECTIVE. Our goal was to describe the variability of sleep duration (time in bed per 24 hours) in healthy children from 1 to 10 years of age in comparison with growth measures.
Activation for newly learned words in left medial-temporal lobe during toddlers' sleep is associated with memory for words
DOI:10.1016/j.cub.2021.09.058 URL [本文引用: 2]
Neuroimaging the sleeping brain: Insight on memory functioning in infants and toddlers
Sleep-dependent selective imitation in infants
Gist extraction and sleep in 12-month-old infants
DOI:10.1016/j.nlm.2016.08.021 URL [本文引用: 2]
Naps promote flexible memory retrieval in 12- month-old infants
DOI:10.1002/dev.v58.7 URL [本文引用: 2]
Sleep spindles in midday naps enhance learning in preschool children
DOI:10.1073/pnas.1306418110
PMID:24062429
[本文引用: 2]
Despite the fact that midday naps are characteristic of early childhood, very little is understood about the structure and function of these sleep bouts. Given that sleep benefits memory in young adults, it is possible that naps serve a similar function for young children. However, children transition from biphasic to monophasic sleep patterns in early childhood, eliminating the nap from their daily sleep schedule. As such, naps may contain mostly light sleep stages and serve little function for learning and memory during this transitional age. Lacking scientific understanding of the function of naps in early childhood, policy makers may eliminate preschool classroom nap opportunities due to increasing curriculum demands. Here we show evidence that classroom naps support learning in preschool children by enhancing memories acquired earlier in the day compared with equivalent intervals spent awake. This nap benefit is greatest for children who nap habitually, regardless of age. Performance losses when nap-deprived are not recovered during subsequent overnight sleep. Physiological recordings of naps support a role of sleep spindles in memory performance. These results suggest that distributed sleep is critical in early learning; when short-term memory stores are limited, memory consolidation must take place frequently.
Sleep and early cortical development
Sleep is increasingly recognized as a key process in neurodevelopment. Animal data show that sleep is essential for the maturation of fundamental brain functions, and growing epidemiological findings indicate that children with early sleep disturbance suffer from later cognitive, attentional, and psychosocial problems. Still, major gaps exist in understanding processes underlying links between sleep and neurodevelopment. One challenge is to translate findings from animal research to humans. In this review, we describe parallels and differences in sleep and development of the cortex in humans and animals and discuss emerging questions.
Overlapping memory replay during sleep builds cognitive schemata
DOI:10.1016/j.tics.2011.06.004
PMID:21764357
[本文引用: 1]
Sleep enhances integration across multiple stimuli, abstraction of general rules, insight into hidden solutions and false memory formation. Newly learned information is better assimilated if compatible with an existing cognitive framework or schema. This article proposes a mechanism by which the reactivation of newly learned memories during sleep could actively underpin both schema formation and the addition of new knowledge to existing schemata. Under this model, the overlapping replay of related memories selectively strengthens shared elements. Repeated reactivation of memories in different combinations progressively builds schematic representations of the relationships between stimuli. We argue that this selective strengthening forms the basis of cognitive abstraction, and explain how it facilitates insight and false memory formation.Copyright © 2011 Elsevier Ltd. All rights reserved.
Slow wave sleep in naps supports episodic memories in early childhood
Relations between sleep patterns early in life and brain development: A review
DOI:10.1016/j.dcn.2022.101130 URL [本文引用: 1]
Sleep and human cognitive development
DOI:10.1016/j.smrv.2021.101472 URL [本文引用: 3]
Neurobiology of REM and NREM sleep
This paper presents an overview of the current knowledge of the neurophysiology and cellular pharmacology of sleep mechanisms. It is written from the perspective that recent years have seen a remarkable development of knowledge about sleep mechanisms, due to the capability of current cellular neurophysiological, pharmacological and molecular techniques to provide focused, detailed, and replicable studies that have enriched and informed the knowledge of sleep phenomenology and pathology derived from electroencephalographic (EEG) analysis. This chapter has a cellular and neurophysiological/neuropharmacological focus, with an emphasis on rapid eye movement (REM) sleep mechanisms and non-REM (NREM) sleep phenomena attributable to adenosine. The survey of neuronal and neurotransmitter-related brainstem mechanisms of REM includes monoamines, acetylcholine, the reticular formation, a new emphasis on GABAergic mechanisms and a discussion of the role of orexin/hypcretin in diurnal consolidation of REM sleep. The focus of the NREM sleep discussion is on the basal forebrain and adenosine as a mediator of homeostatic control. Control is through basal forebrain extracellular adenosine accumulation during wakefulness and inhibition of wakefulness-active neurons. Over longer periods of sleep loss, there is a second mechanism of homeostatic control through transcriptional modification. Adenosine acting at the A1 receptor produces an up-regulation of A1 receptors, which increases inhibition for a given level of adenosine, effectively increasing the gain of the sleep homeostat. This second mechanism likely occurs in widespread cortical areas as well as in the basal forebrain. Finally, the results of a new series of experimental paradigms in rodents to measure the neurocognitive effects of sleep loss and sleep interruption (modeling sleep apnea) provide animal model data congruent with those in humans.
The deferred imitation task as a nonverbal measure of declarative memory
DOI:10.1073/pnas.92.16.7580
PMID:7638234
[本文引用: 1]
We tested amnesic patients, patients with frontal lobe lesions, and control subjects with the deferred imitation task, a nonverbal test used to demonstrate memory abilities in human infants. On day 1, subjects were given sets of objects to obtain a baseline measure of their spontaneous performance of target actions. Then different event sequences were modeled with the object sets. On day 2, the objects were given to the subjects again, first without any instructions to imitate the sequences, and then with explicit instructions to imitate the actions exactly as they had been modeled. Control subjects and frontal lobe patients reproduced the events under both uninstructed and instructed conditions. In contrast, performance by the amnesic patients did not significantly differ from that of a second control group who had the same opportunities to handle the objects but were not shown the modeled actions. These findings suggest that deferred imitation is dependent on the brain structures essential for declarative memory that are damaged in amnesia, and they support the view that infants who imitate actions after long delays have an early capacity for long-term declarative memory.
Memory-related hippocampal activation during sleep and temporal memory in toddlers
DOI:10.1016/j.dcn.2020.100908 URL [本文引用: 2]
Social, motor, and cognitive development through the lens of sleep network dynamics in infants and toddlers between 12 and 30 months of age.
Effects of early and late nocturnal sleep on declarative and procedural memory
DOI:10.1162/jocn.1997.9.4.534
PMID:23968216
[本文引用: 2]
Recall of paired-associate lists (declarative memory) and mirror-tracing skills (procedural memory) was assessed after retention intervals defined over early and late nocturnal sleep. In addition, effects of sleep on recall were compared with those of early and late retention intervals filled with wakefulness. Twenty healthy men served as subjects. Saliva cortisol concentrations were determined before and after the retention intervals to determine pituitary-adrenal secretory activity. Sleep was determined somnopolygraphically. Sleep generally enhanced recall when compared with the effects of corresponding retention intervals of wakefulness. The benefit from sleep on recall depended on the phase of sleep and on the type of memory: Recall of paired-associate lists improved more during early sleep, and recall of mirror-tracing skills improved more during late sleep. The effects may reflect different influences of slow wave sleep (SWS) and rapid eye movement (REM) sleep since time in SWS was 5 times longer during the early than late sleep retention interval, and time in REM sleep was twice as long during late than early sleep (p < 0.005). Changes in cortisol concentrations, which independently of sleep and wakefulness were lower during early retention intervals than late ones, cannot account for the effects of sleep on memory. The experiments for the first time dissociate specific effects of early and late sleep on two principal types of memory, declarative and procedural, in humans.
Memory-related hippocampal activation in the sleeping toddler
DOI:10.1073/pnas.1805572115
PMID:29866845
[本文引用: 2]
Nonhuman research has implicated developmental processes within the hippocampus in the emergence and early development of episodic memory, but methodological challenges have hindered assessments of this possibility in humans. Here, we delivered a previously learned song and a novel song to 2-year-old toddlers during natural nocturnal sleep and, using functional magnetic resonance imaging, found that hippocampal activation was stronger for the learned song compared with the novel song. This was true regardless of whether the song was presented intact or backwards. Toddlers who remembered where and in the presence of which toy character they heard the song exhibited stronger hippocampal activation for the song. The results establish that hippocampal activation in toddlers reflects past experiences, persists despite some alteration of the stimulus, and is associated with behavior. This research sheds light on early hippocampal and memory functioning and offers an approach to interrogate the neural substrates of early memory.
The relationships between self-efficacy, self-control, chronotype, procrastination and sleep problems in young adults
DOI:10.1080/07420528.2019.1607370
PMID:31070062
[本文引用: 1]
The main aim of our study was to examine whether there was a relationship between psychological characteristics such as self-efficacy, self-control and chronotype as well as procrastination on the one hand and sleep problems on the other. There were 315 young adults aged between 18 and 27 years ( = 20.57). We used the General Procrastination Scale, the General Self-Efficacy Scale (GSES), Brief Self-Control Scale, the Composite Scale of Morningness (CSM) and the Pittsburgh Sleep Quality Index (PSQI). Our results indicated that low self-efficacy, low self-control and eveningness were positive predictors of procrastination. The reciprocal relationship exists between procrastination and sleep problems. Procrastination positively contributed to sleep problems, whereas sleep problems were a negative predictor of procrastination.
Effects of a brief afternoon nap on declarative and procedural memory
DOI:10.1016/j.nlm.2022.107662 URL [本文引用: 2]
Odor cues during slow-wave sleep prompt declarative memory consolidation
DOI:10.1126/science.1138581
PMID:17347444
[本文引用: 1]
Sleep facilitates memory consolidation. A widely held model assumes that this is because newly encoded memories undergo covert reactivation during sleep. We cued new memories in humans during sleep by presenting an odor that had been presented as context during prior learning, and so showed that reactivation indeed causes memory consolidation during sleep. Re-exposure to the odor during slow-wave sleep (SWS) improved the retention of hippocampus-dependent declarative memories but not of hippocampus-independent procedural memories. Odor re-exposure was ineffective during rapid eye movement sleep or wakefulness or when the odor had been omitted during prior learning. Concurring with these findings, functional magnetic resonance imaging revealed significant hippocampal activation in response to odor re-exposure during SWS.
Back to basics: Are-evaluation of the relevance of imprinting in the genesis of Bowlby’s attachment theory
DOI:10.3389/fpsyg.2022.1033746
URL
[本文引用: 1]
Attachment theory is one of the key theoretical constructs that underpin explorations of human bonding, taking its current form in John Bowlby’s amalgamation of ideas from psychoanalysis, developmental psychology and ethology. Such a period of interdisciplinary exchange, and Bowlby’s interest in Lorenz’ concept of imprinting in particular, have been subject to rather historical and biographical studies, leaving a fine-grained theoretical scrutiny of the exact relationship between imprinting and attachment still pending. This paper attempts to remedy such an omission by exploring the relationships between these two constructs. It critically reviews the theories of imprinting in general, of human imprinting in particular, and of attachment; analysis of the links between these processes bring to the foreground the distinction between supra-individual vs. individual aspects of bonding, the relevance of ‘proto-attachment’ phases before ‘proper’ Bowlbyan attachment is attained, and the role of communicative signals during such early phases. The paper outlines potential benefits of considering such elements in the study of early social cognition, particularly in respect of the study of the gaze and the infant-directed communicative register.
Slow-wave activity and sigma activities are associated with psychomotor development at 8 months of age.
Sleep-dependent motor memory consolidation in healthy adults: A meta-analysis
DOI:S0149-7634(20)30497-8
PMID:32730847
[本文引用: 3]
It is widely accepted that sleep better facilitates the consolidation of motor memories than does a corresponding wake interval (King et al., 2017). However, no in-depth analysis of the various motor tasks and their relative sleep gain has been conducted so far. Therefore, the present meta-analysis considered 48 studies with a total of 53 sleep (n = 829) and 53 wake (n = 825) groups. An overall comparison between all sleep and wake groups resulted in a small effect for the relative sleep gain in motor memory consolidation (g = 0.43). While no subgroup differences were identified for differing designs, a small effect for the finger tapping task (g = 0.47) and a medium effect for the mirror tracing task (g = 0.62) were found. In summary, the meta-analysis substantiates that sleep generally benefits the consolidation of motor memories. However, to further our understanding of the mechanisms underlying this effect, examining certain task dimensions and their relative sleep gain would be a promising direction for future research.Copyright © 2020 The Author(s). Published by Elsevier Ltd.. All rights reserved.
Timely sleep facilitates declarative memory consolidation in infants
DOI:10.1073/pnas.1414000112
PMID:25583469
[本文引用: 4]
Human infants devote the majority of their time to sleeping. However, very little is known about the role of sleep in early memory processing. Here we test 6- and 12-mo-old infants' declarative memory for novel actions after a 4-h [Experiment (Exp.) 1] and 24-h delay (Exp. 2). Infants in a nap condition took an extended nap (≥30 min) within 4 h after learning, whereas infants in a no-nap condition did not. A comparison with age-matched control groups revealed that after both delays, only infants who had napped after learning remembered the target actions at the test. Additionally, after the 24-h delay, memory performance of infants in the nap condition was significantly higher than that of infants in the no-nap condition. This is the first experimental evidence to our knowledge for an enhancing role of sleep in the consolidation of declarative memories in the first year of life.
Remembering in the context of internal states: The role of sleep for infant memory
DOI:10.1111/cdep.12321
[本文引用: 1]
Research with adults has shown that a person's internal context, or state, influences how memory functions. This factor is rarely considered in research on infant memory, in part because of the practical and ethical difficulties of manipulating these variables in infants. In this article, we argue that models of infant memory will remain limited in scope and accuracy if the internal context of participants is not considered. As a case in point, we present emerging literature on sleep-dependent memory. Our review shows that for infants, timely sleep after a learning experience helps them retain and further process new memories. Studies need to explore the role of prior sleep for encoding, and to tease apart the contributions to infant memory of different types, features, and stages of sleep. We conclude that considering internal states, such as sleep, is necessary for developing a deeper understanding of early human memory.
Sleep confers a benefit for retention of statistical language learning in 6.5 month old infants
DOI:10.1016/j.bandl.2016.05.002 URL [本文引用: 2]
Infants' and toddlers' language, math and socio-emotional development: Evidence for reciprocal relations and differential gender and age effects
DOI:10.3389/fpsyg.2020.580297
URL
[本文引用: 1]
Toddlerhood is characterized by rapid development in several domains, such as language, socio-emotional behavior and emerging math skills all of which are important precursors of school readiness. However, little is known about how these skills develop over time and how they may be interrelated. The current study investigates young children’s development at two time points, with about 7 months in between, assessing their language, socio-emotional and math language and numeracy skills with teacher ratings. The sample includes 577 children from 18 until 36 months of age of 86 childcare classrooms. The results of the autoregressive path analyses showed moderate to strong stability of language, socio-emotional and math language and numeracy skills, although the magnitude of associations was smaller for the latter. The cross-lagged path analyses highlighted the importance of language and socio-emotional skills for development in the other domains. Differential relations were found for the autoregressive and cross-lagged paths depending on gender and age. Language skills appeared a stronger predictor of boys’ socio-emotional and math language and numeracy skill development compared to girls. Girls’ socio-emotional skills predicted growth in math. For boys, socio-emotional and math language and numeracy skills appeared to be unrelated. Language skills showed stronger relations with the development of math language and numeracy skills for younger children as compared to older children. Also, for older children math language and numeracy skills negatively predicted growth in their socio-emotional skills. The findings provide more insights in how language, math language and numeracy skills and socio-emotional skills co-develop in the early years and as such have important implications for interventions aimed to support children’s development.
REM sleep in naps differentially relates to memory consolidation in typical preschoolers and children with Down syndrome
DOI:10.1073/pnas.1811488115
PMID:30373840
[本文引用: 2]
Sleep is recognized as a physiological state associated with learning, with studies showing that knowledge acquisition improves with naps. Little work has examined sleep-dependent learning in people with developmental disorders, for whom sleep quality is often impaired. We examined the effect of natural, in-home naps on word learning in typical young children and children with Down syndrome (DS). Despite similar immediate memory retention, naps benefitted memory performance in typical children but hindered performance in children with DS, who retained less when tested after a nap, but were more accurate after a wake interval. These effects of napping persisted 24 h later in both groups, even after an intervening overnight period of sleep. During naps in typical children, memory retention for object-label associations correlated positively with percent of time in rapid eye movement (REM) sleep. However, in children with DS, a population with reduced REM, learning was impaired, but only after the nap. This finding shows that a nap can increase memory loss in a subpopulation, highlighting that naps are not universally beneficial. Further, in healthy preschooler's naps, processes in REM sleep may benefit learning.
Retrograde amnesia and memory consolidation: A neurobiological perspective
DOI:10.1016/0959-4388(95)80023-9
PMID:7620304
[本文引用: 1]
The fact that information acquired before the onset of amnesia can be lost (retrograde amnesia) has fascinated psychologists, biologists, and clinicians for over 100 years. Studies of retrograde amnesia have led to the concept of memory consolidation, whereby medial temporal lobe structures direct the gradual establishment of memory representations in neocortex. Recent theoretical accounts have inspired a simple neural network model that produces behavior consistent with experimental data and makes these ideas about memory consolidation more concrete. Recent physiological and anatomical findings provide important information about how memory consolidation might actually occur.
Sleep-dependent memory consolidation
DOI:10.1038/nature04286 [本文引用: 3]
Sleep is associated with offline improvement of motor sequence skill in children
Sleep and emotional processing
DOI:S1087-0792(17)30153-3
PMID:29395984
[本文引用: 1]
A growing body of literature suggests that sleep plays a critical role in emotional processing. This review aims at synthesizing current evidence on the role of sleep and sleep loss in the modulation of emotional reactivity, emotional memory formation, empathic behavior, fear conditioning, threat generalization and extinction memory. Behavioral and neurophysiological evidence suggesting that rapid-eye movement (REM) sleep plays an important role in emotional processing is also discussed. Furthermore, we examine the relations between sleep and emotions by reviewing the functional neuroimaging studies that elucidated the brain mechanisms underlying these relations. It is shown that sleep supports the formation of emotional episodic memories throughout all the stages that compose memory processing. On the contrary, sleep loss deteriorates both the encoding of emotional information and the emotional memory consolidation processes. Research is also progressively providing new insights into the protective role of sleep in human emotional homeostasis and regulation, promoting adaptive next-day emotional reactivity. In this respect, evidence converges in indicating that lack of sleep significantly influences emotional reactivity. Moreover, notwithstanding some contradictory findings, the processing of emotionally salient information could mainly benefit from REM sleep. However, some crucial aspects of sleep-dependent emotional modulation remain unclear.Copyright © 2017 Elsevier Ltd. All rights reserved.
Cultural transmission: A view from chimpanzees and human infants
DOI:10.1177/0022022101032002002
URL
[本文引用: 1]
Human beings are biologically adapted for culture in ways that other primates are not, as evidenced most clearly by the fact that only human cultural traditions accumulate modifications over historical time (the ratchet effect). The key adaptation is one that enables individuals to understand other individuals as intentional agents like the self. This species-unique form of social cognition emerges in human ontogeny at around 1 year of age as infants begin to engage with other persons in various kinds of joint attentional activities involving gaze following, social referencing, and gestural communication. Young children’s joint attentional skills then engender some uniquely powerful forms of cultural learning, enabling the acquisition of language, discourse skills, tool use practices, and many other conventional activities. These novel forms of cultural learning allow human beings to pool their cognitive resources both contemporaneously and over historical time in ways that are unique in the animal kingdom.
Sleep spindles as facilitators of memory formation and learning
Memory stabilization with targeted reactivation during human slow-wave sleep
DOI:10.1073/pnas.1201072109
PMID:22691500
[本文引用: 2]
It is believed that neural representations of recent experiences become reactivated during sleep, and that this process serves to stabilize associated memories in long-term memory. Here, we initiated this reactivation process for specific memories during slow-wave sleep. Participants studied 50 object-location associations with object-related sounds presented concurrently. For half of the associations, the related sounds were re-presented during subsequent slow-wave sleep while participants underwent functional MRI. Compared with control sounds, related sounds were associated with increased activation of right parahippocampal cortex. Postsleep memory accuracy was positively correlated with sound-related activation during sleep in various brain regions, including the thalamus, bilateral medial temporal lobe, and cerebellum. In addition, postsleep memory accuracy was also positively correlated with pre- to postsleep changes in parahippocampal-medial prefrontal connectivity during retrieval of reactivated associations. Our results suggest that the brain is differentially activated by studied and unstudied sounds during deep sleep and that the thalamus and medial temporal lobe are involved in establishing the mnemonic consequences of externally triggered reactivation of associative memories.
Associations between gross motor skills and cognitive development in toddlers
DOI:S0378-3782(18)30746-1
PMID:30965194
[本文引用: 1]
The early years of life are critical for motor and cognitive development. A better understanding is needed on the associations between the control and development of motor and cognitive tasks.This study aimed to examine the association between gross motor skills and cognitive development in toddlers.Cross-sectional study.This study included 335 toddlers (aged 19.80 ± 4.08 months, 53.7% boys) from 30 childcare services in Australia.Children were assessed on gross motor skills (Peabody Developmental Motor Scales 2nd Edition; PDMS-2) and cognitive development (Bayley Scales of Infant and Toddler development 3rd edition; Bayley-III).A one-way ANCOVA was conducted to assess associations between gross motor skills and cognitive development controlling for childcare center, sex, age, body mass index and socioeconomic status.The average scores were 96.41 ± 9.84 for gross motor skills (range gross motor quotient 35-165) and 11.45 ± 3.03 for cognitive development (range standard score 1-19). There was a significant positive association between gross motor skills and cognition, F = 12.245, p < 0.001. Both locomotion and object manipulation were significantly positively associated with cognition, F = 14.607, p < 0.001 and F = 9.039, p < 0.001 respectively.Levels of gross motor skills are positively associated with cognitive development in this sample of Australian toddlers. Results reinforce the need for early commencement of gross motor skill promotion as this might be important for cognitive development in the early years.Copyright © 2019 Elsevier B.V. All rights reserved.
A refined model of sleep and the time course of memory formation
DOI:10.1017/S0140525X05000026
URL
[本文引用: 2]
Research in the neurosciences continues to provide evidence that sleep plays a role in the processes of learning and memory. There is less of a consensus, however, regarding the precise stages of memory development during which sleep is considered a requirement, simply favorable, or not important. This article begins with an overview of recent studies regarding sleep and learning, predominantly in the procedural memory domain, and is measured against our current understanding of the mechanisms that govern memory formation. Based on these considerations, I offer a new neurocognitive framework of procedural learning, consisting first of acquisition, followed by two specific stages of consolidation, one involving a process of stabilization, the other involving enhancement, whereby delayed learning occurs. Psychophysiological evidence indicates that initial acquisition does not rely fundamentally on sleep. This also appears to be true for the stabilization phase of consolidation, with durable representations, resistant to interference, clearly developing in a successful manner during time awake (or just time, per se). In contrast, the consolidation stage, resulting in additional/enhanced learning in the absence of further rehearsal, does appear to rely on the process of sleep, with evidence for specific sleep-stage dependencies across the procedural domain. Evaluations at a molecular, cellular, and systems level currently offer several sleep specific candidates that could play a role in sleep-dependent learning. These include the upregulation of select plasticity-associated genes, increased protein synthesis, changes in neurotransmitter concentration, and specific electrical events in neuronal networks that modulate synaptic potentiation.
Practice with sleep makes perfect: Sleep-dependent motor skill learning
DOI:10.1016/s0896-6273(02)00746-8
PMID:12123620
[本文引用: 1]
Improvement in motor skill performance is known to continue for at least 24 hr following training, yet the relative contributions of time spent awake and asleep are unknown. Here we provide evidence that a night of sleep results in a 20% increase in motor speed without loss of accuracy, while an equivalent period of time during wake provides no significant benefit. Furthermore, a significant correlation exists between the improved performance overnight and the amount of stage 2 NREM sleep, particularly late in the night. This finding of sleep-dependent motor skill improvement may have important implications for the efficient learning of all skilled actions in humans.
Overnight therapy? The role of sleep in emotional brain processing
DOI:10.1037/a0016570 URL [本文引用: 1]
'Sleep-dependent' memory consolidation? Brief periods of post-training rest and sleep provide an equivalent benefit for both declarative and procedural memory
Sleep-dependent consolidation of procedural motor memories in children and adults: The pre-sleep level of performance matters
DOI:10.1111/j.1467-7687.2012.01146.x
PMID:22709400
[本文引用: 2]
In striking contrast to adults, in children sleep following training a motor task did not induce the expected (offline) gain in motor skill performance in previous studies. Children normally perform at distinctly lower levels than adults. Moreover, evidence in adults suggests that sleep dependent offline gains in skill essentially depend on the pre-sleep level of performance. Against this background, we asked whether improving children's performance on a motor sequence learning task by extended training to levels approaching those of adults would enable sleep-associated gains in motor skill in this age group also. Children (4-6 years) and adults (18-35 years) performed on the motor sequence learning task (button-box task) before and after ~2-hour retention intervals including either sleep (midday nap) or wakefulness. Whereas one group of children and adults, respectively, received the standard amount of 10 blocks of training before retention intervals of sleep or wakefulness, a further group of children received an extended training on 30 blocks (distributed across 3 days). A further group of adults received a restricted training on only two blocks before the retention intervals. Children after standard training reached lowest performance levels, whereas in adults performance after standard training was highest. Children with extended training and adults after reduced training reached intermediate performance levels. Only at these intermediate performance levels did sleep induce significant gains in motor sequence skill, whereas performance did not benefit from sleep in the low-performing children or in the high-performing adults. Spindle counts in the post-training nap were correlated with performance gains at retrieval only in the adults benefitting from sleep. We conclude that, across age groups, sleep induces the most robust gain in motor skill at an intermediate pre-sleep performance level. In low-performing children sleep-dependent improvements in skill may be revealed only after enhancing the pre-sleep performance level by extended training.© 2012 Blackwell Publishing Ltd.
Goodnight book: Sleep consolidation improves word learning via storybooks
DOI:10.3389/fpsyg.2014.00184
PMID:24624111
[本文引用: 2]
Reading the same storybooks repeatedly helps preschool children learn words. In addition, sleeping shortly after learning also facilitates memory consolidation and aids learning in older children and adults. The current study explored how sleep promotes word learning in preschool children using a shared storybook reading task. Children were either read the same story repeatedly or different stories and either napped after the stories or remained awake. Children's word retention were tested 2.5 h later, 24 h later, and 7 days later. Results demonstrate strong, persistent effects for both repeated readings and sleep consolidation on young children's word learning. A key finding is that children who read different stories before napping learned words as well as children who had the advantage of hearing the same story. In contrast, children who read different stories and remained awake never caught up to their peers on later word learning tests. Implications for educational practices are discussed.
Sleep promotes branch-specific formation of dendritic spines after learning
DOI:10.1126/science.1249098
PMID:24904169
[本文引用: 1]
How sleep helps learning and memory remains unknown. We report in mouse motor cortex that sleep after motor learning promotes the formation of postsynaptic dendritic spines on a subset of branches of individual layer V pyramidal neurons. New spines are formed on different sets of dendritic branches in response to different learning tasks and are protected from being eliminated when multiple tasks are learned. Neurons activated during learning of a motor task are reactivated during subsequent non-rapid eye movement sleep, and disrupting this neuronal reactivation prevents branch-specific spine formation. These findings indicate that sleep has a key role in promoting learning-dependent synapse formation and maintenance on selected dendritic branches, which contribute to memory storage. Copyright © 2014, American Association for the Advancement of Science.
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