Sleep-Dependent Memory Consolidation and Incremental Sentence Comprehension: Computational Dependencies during Language Learning as Revealed by Neuronal Oscillations

We hypothesise a beneficial influence of sleep on the consolidation of the combinatorial mechanisms underlying incremental sentence comprehension. These predictions are grounded in recent work examining the effect of sleep on the consolidation of linguistic information, which demonstrate that sleep-dependent neurophysiological activity consolidates the meaning of novel words and simple grammatical rules. However, the sleep-dependent consolidation of sentence-level combinatorics has not been studied to date. Here, we propose that dissociable aspects of sleep neurophysiology consolidate two different types of combinatory mechanisms in human language: sequence-based (order-sensitive) and dependency-based (order-insensitive) combinatorics. The distinction between the two types of combinatorics is motivated both by cross-linguistic considerations and the neurobiological underpinnings of human language. Unifying this perspective with principles of sleep-dependent memory consolidation, we posit that a function of sleep is to optimise the consolidation of sequence-based knowledge (the when) and the establishment of semantic schemas of unordered items (the what) that underpin cross-linguistic variations in sentence comprehension. This hypothesis builds on the proposal that sleep is involved in the construction of predictive codes, a unified principle of brain function that supports incremental sentence comprehension. Finally, we discuss neurophysiological measures (EEG/MEG) that could be used to test these claims, such as the quantification of neuronal oscillations, which reflect basic mechanisms of information processing in the brain.

[1]  Luc H. Arnal,et al.  A Neurophysiological Perspective on Speech Processing in "The Neurobiology of Language" , 2015 .

[2]  M. L. Lambon Ralph,et al.  The Role of Sleep Spindles and Slow-Wave Activity in Integrating New Information in Semantic Memory , 2013, The Journal of Neuroscience.

[3]  T. Harmony The functional significance of delta oscillations in cognitive processing , 2013, Front. Integr. Neurosci..

[4]  E. A. Graves The effect of sleep upon retention , 1936 .

[5]  A. Friederici,et al.  Neural basis of processing sequential and hierarchical syntactic structures , 2007, Human brain mapping.

[6]  D. Poeppel,et al.  Cortical Tracking of Hierarchical Linguistic Structures in Connected Speech , 2015, Nature Neuroscience.

[7]  C. Degueldre,et al.  Concurrent Synaptic and Systems Memory Consolidation during Sleep , 2013, The Journal of Neuroscience.

[8]  Krish D. Singh,et al.  Visual gamma oscillations: The effects of stimulus type, visual field coverage and stimulus motion on MEG and EEG recordings , 2013, NeuroImage.

[9]  A. Staub,et al.  The Timecourse of Sentence Processing in the Brain , 2016 .

[10]  M. Bastiaansen,et al.  A predictive coding framework for rapid neural dynamics during sentence-level language comprehension , 2015, Cortex.

[11]  G. Tononi,et al.  Non-fluent aphasia and neural reorganization after speech therapy: insights from human sleep electrophysiology and functional magnetic resonance imaging. , 2010, Archives italiennes de biologie.

[12]  J. Born,et al.  In search of a role of REM sleep in memory formation , 2015, Neurobiology of Learning and Memory.

[13]  Manuel Schabus,et al.  Sleep spindles and their significance for declarative memory consolidation. , 2004, Sleep.

[14]  Olaf Sporns,et al.  Complex network measures of brain connectivity: Uses and interpretations , 2010, NeuroImage.

[15]  Diane M. Beck,et al.  Pulsed Out of Awareness: EEG Alpha Oscillations Represent a Pulsed-Inhibition of Ongoing Cortical Processing , 2011, Front. Psychology.

[16]  Sarah Brown-Schmidt,et al.  The hippocampus and the flexible use and processing of language , 2012, Front. Hum. Neurosci..

[17]  E. Whitham,et al.  Scalp electrical recording during paralysis: Quantitative evidence that EEG frequencies above 20Hz are contaminated by EMG , 2007, Clinical Neurophysiology.

[18]  J. Hobson,et al.  Not only…but also: REM sleep creates and NREM Stage 2 instantiates landmark junctions in cortical memory networks , 2015, Neurobiology of Learning and Memory.

[19]  Ole Jensen,et al.  Sleep Promotes the Extraction of Grammatical Rules , 2013, PloS one.

[20]  Bertram Opitz,et al.  Interactions of the hippocampal system and the prefrontal cortex in learning language-like rules , 2003, NeuroImage.

[21]  James W. Minett,et al.  Delta, theta, beta, and gamma brain oscillations index levels of auditory sentence processing , 2016, NeuroImage.

[22]  W. Klimesch,et al.  EEG alpha oscillations: The inhibition–timing hypothesis , 2007, Brain Research Reviews.

[23]  J. Born,et al.  Auditory Closed-Loop Stimulation of the Sleep Slow Oscillation Enhances Memory , 2013, Neuron.

[24]  Uwe Friese,et al.  Theta–gamma coupling during episodic retrieval in the human EEG , 2014, Brain Research.

[25]  E. Bates,et al.  Psycholinguistics: a cross-language perspective. , 2001, Annual review of psychology.

[26]  M. Gaskell,et al.  Sleep-Associated Changes in the Mental Representation of Spoken Words , 2007, Psychological science.

[27]  David Kemmerer,et al.  The Cross-Linguistic Prevalence of SOV and SVO Word Orders Reflects the Sequential and Hierarchical Representation of Action in Broca's Area , 2012, Lang. Linguistics Compass.

[28]  B. MacWhinney,et al.  Cue validity and sentence interpretation in English, German, and Italian , 1984 .

[29]  Karsten Steinhauer,et al.  Brain signatures of artificial language processing: Evidence challenging the critical period hypothesis , 2002, Proceedings of the National Academy of Sciences of the United States of America.

[30]  Matthias Schlesewsky,et al.  The resolution of case conflicts from a neurophysiological perspective. , 2005, Brain research. Cognitive brain research.

[31]  Marina Schmid,et al.  An Introduction To The Event Related Potential Technique , 2016 .

[32]  James M. McQueen,et al.  Interaction between episodic and semantic memory networks in the acquisition and consolidation of novel spoken words , 2017, Brain and Language.

[33]  J. Fell,et al.  The role of phase synchronization in memory processes , 2011, Nature Reviews Neuroscience.

[34]  Stuart M. Fogel,et al.  Dissociable learning-dependent changes in REM and non-REM sleep in declarative and procedural memory systems , 2007, Behavioural Brain Research.

[35]  Rebecca Frost,et al.  Sleep Underpins the Plasticity of Language Production , 2014, Psychological science.

[36]  Ina Bornkessel-Schlesewsky,et al.  Neural mechanisms of sentence comprehension based on predictive processes and decision certainty: Electrophysiological evidence from non-canonical linearizations in a flexible word order language , 2016, Brain Research.

[37]  James M. McQueen,et al.  Changes in Theta and Beta Oscillations as Signatures of Novel Word Consolidation , 2015, Journal of Cognitive Neuroscience.

[38]  Karl J. Friston The free-energy principle: a unified brain theory? , 2010, Nature Reviews Neuroscience.

[39]  Steven C. Pan,et al.  Time for considering the possibility that sleep plays no unique role in motor memory consolidation: Reply to Adi-Japha and Karni (2016). , 2017, Psychological bulletin.

[40]  Sara C. Mednick,et al.  Pharmacologically Increasing Sleep Spindles Enhances Recognition for Negative and High-arousal Memories , 2013, Journal of Cognitive Neuroscience.

[41]  S. Frisch,et al.  The N400 reflects problems of thematic hierarchizing , 2001, Neuroreport.

[42]  M. Bozkurt,et al.  Functional anatomy. , 1980, Equine veterinary journal.

[43]  G. Tononi,et al.  Sleep and the Price of Plasticity: From Synaptic and Cellular Homeostasis to Memory Consolidation and Integration , 2014, Neuron.

[44]  H. Neville,et al.  An Event-Related fMRI Study of Syntactic and Semantic Violations , 2001, Journal of psycholinguistic research.

[45]  Georg Gruber,et al.  Oscillatory Theta Activity during Memory Formation and Its Impact on Overnight Consolidation: A Missing Link? , 2015, Journal of Cognitive Neuroscience.

[46]  Luc H. Arnal,et al.  Transitions in neural oscillations reflect prediction errors generated in audiovisual speech , 2011, Nature Neuroscience.

[47]  Peter Hagoort,et al.  Neural activity during sentence processing as reflected in theta, alpha, beta, and gamma oscillations , 2016, NeuroImage.

[48]  Paul J. Reber,et al.  Sleep facilitates learning a new linguistic rule , 2014, Neuropsychologia.

[49]  M. Schlesewsky,et al.  The N400 as a correlate of interpretively relevant linguistic rules: Evidence from Hindi , 2009, Neuropsychologia.

[50]  B. Staresina,et al.  Oscillations and Episodic Memory: Addressing the Synchronization/Desynchronization Conundrum , 2016, Trends in Neurosciences.

[51]  A. Graybiel,et al.  Activity of striatal neurons reflects dynamic encoding and recoding of procedural memories , 2005, Nature.

[52]  G. Tononi,et al.  Sleep function and synaptic homeostasis. , 2006, Sleep medicine reviews.

[53]  N. Burgess,et al.  Brain oscillations and memory , 2010, Current Opinion in Neurobiology.

[54]  Robert Stickgold,et al.  Sleep Spindle Activity is Associated with the Integration of New Memories and Existing Knowledge , 2010, The Journal of Neuroscience.

[55]  E. Basar,et al.  Links of Consciousness, Perception, and Memory by Means of Delta Oscillations of Brain , 2016, Front. Psychol..

[56]  Sean M Montgomery,et al.  Entrainment of Neocortical Neurons and Gamma Oscillations by the Hippocampal Theta Rhythm , 2008, Neuron.

[57]  Jutta L. Mueller,et al.  Neural networks involved in learning lexical-semantic and syntactic information in a second language , 2014, Front. Psychol..

[58]  Björn Rasch,et al.  Cueing vocabulary during sleep increases theta activity during later recognition testing. , 2015, Psychophysiology.

[59]  J. Martinerie,et al.  The brainweb: Phase synchronization and large-scale integration , 2001, Nature Reviews Neuroscience.

[60]  Morten H. Christiansen,et al.  Arbitrariness, Iconicity, and Systematicity in Language , 2015, Trends in Cognitive Sciences.

[61]  D. Perani,et al.  Dorsal and ventral pathways in language development , 2013, Brain and Language.

[62]  B. Staresina,et al.  Hierarchical nesting of slow oscillations, spindles and ripples in the human hippocampus during sleep , 2015, Nature Neuroscience.

[63]  R. Knight,et al.  The functional role of cross-frequency coupling , 2010, Trends in Cognitive Sciences.

[64]  Sabine Weiss,et al.  “Too Many betas do not Spoil the Broth”: The Role of Beta Brain Oscillations in Language Processing , 2012, Front. Psychology.

[65]  D. Cohen,et al.  Comparison of the magnetoencephalogram and electroencephalogram. , 1979, Electroencephalography and clinical neurophysiology.

[66]  Matthew H. Davis,et al.  A complementary systems account of word learning: neural and behavioural evidence , 2009, Philosophical Transactions of the Royal Society B: Biological Sciences.

[67]  D. Vernon,et al.  Interpreting EEG alpha activity , 2014, Neuroscience & Biobehavioral Reviews.

[68]  Kara Morgan-Short,et al.  Explicit and Implicit Second Language Training Differentially Affect the Achievement of Native-like Brain Activation Patterns , 2012, Journal of Cognitive Neuroscience.

[69]  F. Castellanos,et al.  Entrainment of neural oscillations as a modifiable substrate of attention , 2014, Trends in Cognitive Sciences.

[70]  J. Lisman,et al.  The Theta-Gamma Neural Code , 2013, Neuron.

[71]  John C J Hoeks,et al.  Seeing words in context: the interaction of lexical and sentence level information during reading. , 2004, Brain research. Cognitive brain research.

[72]  M. Ullman A neurocognitive perspective on language: The declarative/procedural model , 2001, Nature Reviews Neuroscience.

[73]  Isabel C. Hutchison,et al.  The role of REM sleep theta activity in emotional memory , 2015, Front. Psychol..

[74]  K. McGregor,et al.  Sleep Disorders as a Risk to Language Learning and Use. , 2015, EBP briefs.

[75]  Annette Sterr,et al.  The reorganisation of memory during sleep. , 2014, Sleep medicine reviews.

[76]  Jan Born,et al.  A Role of Sleep in Forming Predictive Codes , 2017 .

[77]  A. Kleinschmidt,et al.  Brain Networks and α-Oscillations: Structural and Functional Foundations of Cognitive Control , 2016, Trends in Cognitive Sciences.

[78]  Dorothy Tse,et al.  References and Notes Supporting Online Material Materials and Methods Figs. S1 to S5 Tables S1 to S3 Electron Impact (ei) Mass Spectra Chemical Ionization (ci) Mass Spectra References Schemas and Memory Consolidation Research Articles Research Articles Research Articles Research Articles , 2022 .

[79]  Karl J. Friston,et al.  The Functional Anatomy of Time: What and When in the Brain , 2016, Trends in Cognitive Sciences.

[80]  G. Tononi,et al.  Time to Be SHY? Some Comments on Sleep and Synaptic Homeostasis , 2012, Neural plasticity.

[81]  James L. McClelland,et al.  What Learning Systems do Intelligent Agents Need? Complementary Learning Systems Theory Updated , 2016, Trends in Cognitive Sciences.

[82]  B. Rasch,et al.  The beneficial role of memory reactivation for language learning during sleep: A review , 2017, Brain and Language.

[83]  Ina Bornkessel-Schlesewsky,et al.  The Role of Prominence Information in the Real-Time Comprehension of Transitive Constructions: A Cross-Linguistic Approach , 2009, Lang. Linguistics Compass.

[84]  Rajesh P. N. Rao,et al.  Predictive Coding , 2019, A Blueprint for the Hard Problem of Consciousness.

[85]  Asaf Gilboa,et al.  Neurobiology of Schemas and Schema-Mediated Memory , 2017, Trends in Cognitive Sciences.

[86]  G. Tononi,et al.  Sleep and synaptic renormalization: a computational study. , 2010, Journal of neurophysiology.

[87]  J. Rauschecker,et al.  Neurobiological roots of language in primate audition: common computational properties , 2015, Trends in Cognitive Sciences.

[88]  Robert Stickgold,et al.  Cerebral Cortex doi:10.1093/cercor/bhi043 The Functional Anatomy of Sleep-dependent Visual Skill Learning , 2005 .

[89]  György Buzsáki,et al.  What does gamma coherence tell us about inter-regional neural communication? , 2015, Nature Neuroscience.

[90]  Matthias Schlesewsky,et al.  The extended argument dependency model: a neurocognitive approach to sentence comprehension across languages. , 2006, Psychological review.

[91]  Balthasar Bickel,et al.  Language evolution: syntax before phonology? , 2014, Proceedings of the Royal Society B: Biological Sciences.

[92]  B. Rasch,et al.  Boosting Vocabulary Learning by Verbal Cueing During Sleep. , 2015, Cerebral cortex.

[93]  Kevin R Peters,et al.  Changes in the density of stage 2 sleep spindles following motor learning in young and older adults , 2008, Journal of sleep research.

[94]  P. Enticott,et al.  Transcranial electrical stimulation during sleep enhances declarative (but not procedural) memory consolidation: Evidence from a meta-analysis , 2016, Neuroscience & Biobehavioral Reviews.

[95]  G Buzsáki,et al.  The hippocampo-neocortical dialogue. , 1996, Cerebral cortex.

[96]  P. Lewis,et al.  Schema-conformant memories are preferentially consolidated during REM sleep , 2015, Neurobiology of Learning and Memory.

[97]  Angela D Friederici,et al.  ERP evidence for different strategies in the processing of case markers in native speakers and non-native learners , 2007, BMC Neuroscience.

[98]  A. Friederici,et al.  Brain Correlates of Language Learning: The Neuronal Dissociation of Rule-Based versus Similarity-Based Learning , 2004, The Journal of Neuroscience.

[99]  Jack J. Lin,et al.  Direct brain recordings reveal hippocampal rhythm underpinnings of language processing , 2016, Proceedings of the National Academy of Sciences.

[100]  David Poeppel,et al.  Cortical oscillations and speech processing: emerging computational principles and operations , 2012, Nature Neuroscience.

[101]  A. Giuditta Sleep memory processing: the sequential hypothesis , 2014, Front. Syst. Neurosci..

[102]  M. Bastiaansen,et al.  Fast oscillatory dynamics during language comprehension: Unification versus maintenance and prediction? , 2015, Brain and Language.

[103]  R. Oostenveld,et al.  Theta and Gamma Oscillations Predict Encoding and Retrieval of Declarative Memory , 2006, The Journal of Neuroscience.

[104]  M. Gareth Gaskell,et al.  Does Sleep Improve Your Grammar? Preferential Consolidation of Arbitrary Components of New Linguistic Knowledge , 2016, PloS one.

[105]  Simon Hanslmayr,et al.  Brain oscillatory subsequent memory effects differ in power and long-range synchronization between semantic and survival processing , 2013, NeuroImage.

[106]  P. Lewis,et al.  Overnight consolidation aids the transfer of statistical knowledge from the medial temporal lobe to the striatum. , 2013, Cerebral cortex.

[107]  Gernot G. Supp,et al.  Spindle activity phase-locked to sleep slow oscillations , 2016, NeuroImage.

[108]  W. Klimesch EEG alpha and theta oscillations reflect cognitive and memory performance: a review and analysis , 1999, Brain Research Reviews.

[109]  P. Lewis,et al.  Overlapping memory replay during sleep builds cognitive schemata , 2011, Trends in Cognitive Sciences.

[110]  M. Schlesewsky,et al.  Think globally: Cross-linguistic variation in electrophysiological activity during sentence comprehension , 2011, Brain and Language.

[111]  Anne-Catherine Bachoud-Lévi,et al.  Different Neurophysiological Mechanisms Underlying Word and Rule Extraction from Speech , 2007, PloS one.

[112]  G. Rauchs,et al.  The relationships between memory systems and sleep stages , 2005, Journal of sleep research.

[113]  Peter Ford Dominey,et al.  Neural network processing of natural language: II. Towards a unified model of corticostriatal function in learning sentence comprehension and non-linguistic sequencing , 2009, Brain and Language.

[114]  I. Wilhelm,et al.  System consolidation of memory during sleep , 2011, Psychological Research.

[115]  David Poeppel,et al.  Acoustic landmarks drive delta–theta oscillations to enable speech comprehension by facilitating perceptual parsing , 2014, NeuroImage.

[116]  Simon Hanslmayr,et al.  How brain oscillations form memories — A processing based perspective on oscillatory subsequent memory effects , 2014, NeuroImage.

[117]  Sven Bestmann,et al.  Transcranial electrical stimulation , 2017, Current Biology.

[118]  G. Albouy,et al.  Hippocampus and striatum: Dynamics and interaction during acquisition and sleep‐related motor sequence memory consolidation , 2013, Hippocampus.

[119]  Anne-Lise Giraud,et al.  The contribution of frequency-specific activity to hierarchical information processing in the human auditory cortex , 2014, Nature Communications.

[120]  C. N. Cofer Origins of the journal of verbal learning and verbal behavior , 1978 .

[121]  Bruce L. Miller,et al.  What Role Does the Anterior Temporal Lobe Play in Sentence-level Processing? Neural Correlates of Syntactic Processing in Semantic Variant Primary Progressive Aphasia , 2014, Journal of Cognitive Neuroscience.

[122]  Denise M Werchan,et al.  Sleep confers a benefit for retention of statistical language learning in 6.5month old infants , 2017, Brain and Language.

[123]  S. Cole,et al.  Brain Oscillations and the Importance of Waveform Shape , 2017, Trends in Cognitive Sciences.

[124]  Steven C. Pan,et al.  Sleep and motor learning: Is there room for consolidation? , 2015, Psychological bulletin.

[125]  F. Bartlett,et al.  Remembering: A Study in Experimental and Social Psychology , 1932 .

[126]  M. Ullman Contributions of memory circuits to language: the declarative/procedural model , 2004, Cognition.

[127]  D. Bendor,et al.  Does the Hippocampus Map Out the Future? , 2016, Trends in Cognitive Sciences.

[128]  I. Fried,et al.  Sleep Spindles in Humans: Insights from Intracranial EEG and Unit Recordings , 2011, The Journal of Neuroscience.

[129]  Karl J. Friston,et al.  Waking and dreaming consciousness: Neurobiological and functional considerations , 2012, Progress in Neurobiology.

[130]  J. Rauschecker,et al.  Maps and streams in the auditory cortex: nonhuman primates illuminate human speech processing , 2009, Nature Neuroscience.

[131]  Melissa C. Duff,et al.  Expanding the Language Network: Direct Contributions from the Hippocampus , 2016, Trends in Cognitive Sciences.

[132]  Ian Schofield,et al.  A Neocortical Delta Rhythm Facilitates Reciprocal Interlaminar Interactions via Nested Theta Rhythms , 2013, The Journal of Neuroscience.

[133]  Luc H. Arnal Predicting “When” Using the Motor System’s Beta-Band Oscillations , 2012, Front. Hum. Neurosci..

[134]  Reinhold Kliegl,et al.  Cue validity and sentence interpretation in , 2008 .

[135]  M. Schlesewsky,et al.  Reconciling time, space and function: A new dorsal–ventral stream model of sentence comprehension , 2013, Brain and Language.

[136]  R. Stickgold,et al.  REM sleep enhancement of probabilistic classification learning is sensitive to subsequent interference , 2015, Neurobiology of Learning and Memory.

[137]  R. Vertes,et al.  The case against memory consolidation in REM sleep. , 2000, The Behavioral and brain sciences.

[138]  Morten H. Christiansen,et al.  fMRI Syntactic and Lexical Repetition Effects Reveal the Initial Stages of Learning a New Language. , 2016, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[139]  G. Tononi,et al.  Plastic Changes Following Imitation-Based Speech and Language Therapy for Aphasia , 2014, Neurorehabilitation and neural repair.

[140]  M. Ullman The Declarative/Procedural Model , 2020, Theories in Second Language Acquisition.

[141]  Nora A. Herweg,et al.  Theta-Alpha Oscillations Bind the Hippocampus, Prefrontal Cortex, and Striatum during Recollection: Evidence from Simultaneous EEG–fMRI , 2016, The Journal of Neuroscience.

[142]  C. Schroeder,et al.  Low-frequency neuronal oscillations as instruments of sensory selection , 2009, Trends in Neurosciences.

[143]  Jutta L. Mueller L2 in a Nutshell: The Investigation of Second Language Processing in the Miniature Language Model. , 2006 .

[144]  György Buzsáki,et al.  Neural Syntax: Cell Assemblies, Synapsembles, and Readers , 2010, Neuron.

[145]  James L. McClelland Incorporating rapid neocortical learning of new schema-consistent information into complementary learning systems theory. , 2013, Journal of experimental psychology. General.

[146]  Björn Rasch,et al.  Sleep and language learning , 2017, Brain and Language.

[147]  W. Klimesch Alpha-band oscillations, attention, and controlled access to stored information , 2012, Trends in Cognitive Sciences.

[148]  C. Smith Sleep states and memory processes in humans: procedural versus declarative memory systems. , 2001, Sleep medicine reviews.

[149]  Jan-Mathijs Schoffelen,et al.  A Predictive Coding Perspective on Beta Oscillations during Sentence-Level Language Comprehension , 2016, Front. Hum. Neurosci..

[150]  J. Born,et al.  The memory function of sleep , 2010, Nature Reviews Neuroscience.

[151]  Penelope A. Lewis,et al.  Cross-modal transfer of statistical information benefits from sleep , 2016, Cortex.

[152]  A. Engel,et al.  Beta-band oscillations—signalling the status quo? , 2010, Current Opinion in Neurobiology.

[153]  J. Born,et al.  About sleep's role in memory. , 2013, Physiological reviews.

[154]  Jessica D. Payne,et al.  Napping and the selective consolidation of negative aspects of scenes. , 2015, Emotion.

[155]  O. Jensen,et al.  Beyond ERPs: Oscillatory neuronal dynamics , 2012 .

[156]  Jens G. Klinzing,et al.  Sleep’s role in the reconsolidation of declarative memories , 2016, Neurobiology of Learning and Memory.

[157]  M. Pickering,et al.  An integrated theory of language production and comprehension. , 2013, The Behavioral and brain sciences.

[158]  Christopher K. Kovach,et al.  Manifestation of ocular-muscle EMG contamination in human intracranial recordings , 2011, NeuroImage.

[159]  I. Ajzen Nature and operation of attitudes. , 2001, Annual review of psychology.

[160]  K. Paller,et al.  Sleep-based memory processing facilitates grammatical generalization: Evidence from targeted memory reactivation , 2017, Brain and Language.