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Hippocampus Medial Temporal Cortex Itamar Lerner & Mark A. Gluck 1 Center for Molecular & Behavioral Neuroscience, Rutgers University, Newark, NJ, USA Sleep Mediation of Episodic Memory and Associative Sleep Mediation of Episodic Memory and Associative Learning II: Learning II: A Potential Computational Synthesis A Potential Computational Synthesis Background - I Sleep improves performance: Representative findings Qualitative Traits of the Model Evidence from the last decade shows that sleep has an important role in learning and memory. specifically, sleep – and especially Slow-Wave Sleep (SWS) and, sometimes, Rapid-Eye-Movement sleep (REM) – has been shown to improve episodic memory, gist extraction, and rule extrapolation and insight. In addition, it has been shown that following sleep (especially SWS) synaptic strength within cortical and hippocampal circuits is generally decreased, these two findings have often been taken to support different and even contradicting theories about the role of sleep in learning and memory. The current work in progress is a computational approach that seeks to combine a broad range of empirical findings within a uniform neuro-computational framework. Introduction Conclusions Acknowledgements Backhaus J, Born J, Hoeckesfeld R, Fokuhl S, Hohagen F, & Junghanns K (2007). Midlife decline in declarative memory consolidation is correlated with a decline in slow wave sleep. Learning & Memory, 14, 336-341. Ellenbogen JM, Hulbert JC, Stickgold R, Dinges DF, Thompson- Schill SL (2006). Interfering with theories of sleep and memory: sleep, declarative memory, and associative interference. Current Biology, 16, 1290-1294. Gluck MA, Myers CE (1993). Hippocampal mediation of stimulus representation: a computational theory. Hippocampus, 3, 491-516. Lau H, Alger SE, Fishbein W (2011). Relational memory, a daytime naps facilitates abstraction of general concepts. PLoS One, 6. e27139. Liu ZW, Faraguna U, Cirelli C, Tononi G, Gao XB (2010). Direct evidence for wake-related increases and sleep-related decreases in synaptic strength in rodent cortex. Journal of Neuroscience, 30, 8671–8675. Moustafa AA, Myers CE, Gluck MA (2009). A neurocomputational model of classical conditioning phenomena: a putative role for the hippocampal region in associative learning. Brain Research, 1276, 180–195. Plihal W, Born J (1997) Effects of early and late nocturnal sleep on declarative and procedural memory Journal of Cognitive Neuroscience, 9, 534–547. Vyazovskiy VV, Cirelli C, Pfister-Genskow C, Faraguna U, Tononi G (2008). Molecular and electrophysiological evidence for net synaptic potentiation in wake and depression in sleep. Nature Neuroscience, 11, 2, 200–208. References Performance I. Episodic memory: Paired associates learning Dog - Pianist Hole - Sky Memor izing Diamond - Letter Testi ng II. Gist extraction: Learning meaning of Chinese characters: Training : Memorizing relations between the item pairs a,b,c,d,e: a>b, b>c, c>d, d>e, e>f (pairs contain implicit hierarchy: a>b>c>d>e>f) Testing : Hierarchies with 1° separation: b>d, d>e Hierarchies with 2° separation: b>e III. Rule extrapolation: Learning Implicit hierarchy between stimuli General design: training wake/sleep testing Background - II Synaptic strength is reduced during sleep: Representative findings Slope and Amplitude of Excitatory Post-Synaptic Potentials (EPSPs) in the prefrontal cortex of rats decrease following sleep compared to a Sleep-deprivation period. (W – Wake; S – Sleep; Vyazovskiy et al., 2008) Model Principles Differentiation: Representations with a small degree of correlations become largely uncorrelated Unification: Representations that are very correlated to each other are unified to become a single representation. 3. Both of these changes are carried out by deletion of synapses: Differentiation is achieved by deletion of synapses that support activation of neurons common to several representations (thus causing these representations to become uncorrelated). Unification is achieved by deletion of synapses that support activation of neurons that are unique Recall performance increases due to SWS between training and testing Paired-associates task- design. Observation of pairs to be memorized is followed by 12hours of wake or sleep, after which cued recall is tested. A shared pattern in Chinese characters is recognized better after sleep Hierarchy rule is more easily recognized after sleep compared to wake al., 2009) we assert that storing episodic memories, extracting gist information, or extrapolating a classification rule, all crucially depend on gradual learning of stimulus- stimulus associations in the hippocampus during wake. Only after learning these statistical regularities, can the system (Medial Temporal Cortex and Striatum) process appropriate responses. Parsimonious representations facilitate cognitive performance Before sleep: After sleep: Paired- associat es: Gist extracti on: Complete the test sample with activation based on the correct learned pattern Which of the two test samples fit better to the learned patterns? Learned patterns Test sample Learned patterns Test sample Learned patterns Test samples Learned patterns Test samples Objective : Rule learning: Learned structures Test sample Learned structures Test sample To which of the two learned structures does the test sample fit? Differentiation and Unification processes. Each row represents a different memory pattern learned by the hippocampus during wake. Each circle represents a unit (neuron). Red circles - active units; White units – inactive. A. Differentiation B. Unification Supported by Grant #7367437 for “Long- term Mobile Monitoring and Analysis of Sleep-Cognition Relationship” from the National Science Foundation's Smart Health and Wellbeing program to M.A.G. Synaptic deletion during sleep may play a computational role in improving cognitive performance by differentiating and unifying representations 2. Sleep (especially SWS) provides an additional processing stage to the hippocampal representations that were acquired during wake, allowing them to become more parsimonious and consequently boost performance in the subsequent testing phase. This additional stage is based on two processes: 1. Based on our previous NSF-supported modeling (Gluck and Myers, 1993; Moustafa et Ellenboge n et al., 2006 Plihal & Born, 1997 Lau et al., 2011 Backhaus et al., 2011 Changes in cortical Local Field Potential (LFP) in rats in response to stimulation after a period of wake (Sleep Deprived - SD) compared to sleep (Liu et al., 2010) Sleep Gradual learning during wake Sleep extends pattern differentiation Sleep sharpens hippocampal input- to-output correlational differences After sleep-dependent unification and differentiation, each objective is more readily accessed: Input correlation Output correlation Contact == Itamar Lerner, [email protected] Mark Gluck, [email protected]

HippocampusMedial Temporal Cortex Itamar Lerner & Mark A. Gluck 1 Center for Molecular & Behavioral Neuroscience, Rutgers University, Newark, NJ, USA Sleep

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Page 1: HippocampusMedial Temporal Cortex Itamar Lerner & Mark A. Gluck 1 Center for Molecular & Behavioral Neuroscience, Rutgers University, Newark, NJ, USA Sleep

HippocampusMedial Temporal Cortex

Itamar Lerner & Mark A. Gluck1Center for Molecular & Behavioral Neuroscience, Rutgers University, Newark, NJ, USA

Sleep Mediation of Episodic Memory and Associative Learning II:Sleep Mediation of Episodic Memory and Associative Learning II:A Potential Computational SynthesisA Potential Computational Synthesis

Background - I

Sleep improves performance: Representative findings

Qualitative Traits of the Model

Evidence from the last decade shows that sleep has an important role in learning and memory. specifically, sleep – and especially Slow-Wave Sleep (SWS) and, sometimes, Rapid-Eye-Movement sleep (REM) – has been shown to improve episodic memory, gist extraction, and rule extrapolation and insight. In addition, it has been shown that following sleep (especially SWS) synaptic strength within cortical and hippocampal circuits is generally decreased, these two findings have often been taken to support different and even contradicting theories about the role of sleep in learning and memory. The current work in progress is a computational approach that seeks to combine a broad range of empirical findings within a uniform neuro-computational framework.

Introduction

Conclusions

Acknowledgements

Backhaus J, Born J, Hoeckesfeld R, Fokuhl S, Hohagen F, & Junghanns K (2007). Midlife decline in declarative memory consolidation is correlated with a decline in slow wave sleep. Learning & Memory, 14, 336-341.

Ellenbogen JM, Hulbert JC, Stickgold R, Dinges DF, Thompson- Schill SL (2006). Interfering with theories of sleep and memory: sleep, declarative memory, and associative interference. Current Biology, 16, 1290-1294.

Gluck MA, Myers CE (1993). Hippocampal mediation of stimulus representation: a computational theory. Hippocampus, 3, 491-516. Lau H, Alger SE, Fishbein W (2011). Relational memory, a daytime naps facilitates abstraction of general concepts. PLoS One, 6. e27139. Liu ZW, Faraguna U, Cirelli C, Tononi G, Gao XB (2010). Direct evidence for wake-related increases and sleep-related decreases in

synaptic strength in rodent cortex. Journal of Neuroscience, 30, 8671–8675.Moustafa AA, Myers CE, Gluck MA (2009). A neurocomputational model of classical conditioning phenomena: a putative role for the

hippocampal region in associative learning. Brain Research, 1276, 180–195.Plihal W, Born J (1997) Effects of early and late nocturnal sleep on declarative and procedural memory Journal of Cognitive

Neuroscience, 9, 534–547. Vyazovskiy VV, Cirelli C, Pfister-Genskow C, Faraguna U, Tononi G (2008). Molecular and electrophysiological evidence for net

synaptic potentiation in wake and depression in sleep. Nature Neuroscience, 11, 2, 200–208.

References

Per

form

ance

I. Episodic memory: Paired associates learning

Dog - Pianist

Hole - Sky

Memorizing

Diamond - Letter

Testing

II. Gist extraction: Learning meaning of Chinese characters:

Training: Memorizing relations between the item pairs a,b,c,d,e:a>b, b>c, c>d, d>e, e>f (pairs contain implicit hierarchy: a>b>c>d>e>f)

Testing: Hierarchies with 1° separation: b>d, d>eHierarchies with 2° separation: b>e

III. Rule extrapolation: Learning Implicit hierarchy between stimuli

General design: training wake/sleep testing

Background - II

Synaptic strength is reduced during sleep: Representative findings

Slope and Amplitude of Excitatory Post-Synaptic Potentials (EPSPs) in the prefrontal cortex of rats decrease following sleep compared to a Sleep-deprivation period. (W – Wake; S – Sleep; Vyazovskiy et al., 2008)

Model Principles

• Differentiation: Representations with a small degree of correlations become largely uncorrelated

• Unification: Representations that are very correlated to each other are unified to become a single representation.

3. Both of these changes are carried out by deletion of synapses: Differentiation is achieved by deletion of synapses that support activation of neurons common to several representations (thus causing these representations to become uncorrelated). Unification is achieved by deletion of synapses that support activation of neurons that are unique to each representation (thus allowing only neurons common to all these representations to survive, turning these separate representations into a single representation).

Recall performance increases due to SWS between training and testing

Paired-associates task-design. Observation of pairs to be memorized is followed by 12hours of wake or sleep, after which cued recall is tested.

A shared pattern in Chinese characters is recognized better after sleep

Hierarchy rule is more easily recognized after sleep compared to wake

al., 2009) we assert that storing episodic memories, extracting gist information, or extrapolating a classification rule, all crucially depend on gradual learning of stimulus-stimulus associations in the hippocampus during wake. Only after learning these statistical regularities, can the system (Medial Temporal Cortex and Striatum) process appropriate responses.

Parsimonious representations facilitate cognitive performance

Before sleep: After sleep:

Paired-associates:

Gist extraction:

Complete the test sample with activation based on the correct learned pattern

Which of the two test samples fit better to the learned patterns?

Learnedpatterns

Testsample

Learnedpatterns

Testsample

Learnedpatterns Test

samples

Learnedpatterns Test

samples

Objective:

Rule learning:

Learnedstructures

Testsample

Learnedstructures

Testsample

To which of the two learned structures does the test sample fit?

Differentiation and Unification processes. Each row represents a different memory pattern learned by the hippocampus during wake. Each circle represents a unit (neuron). Red circles - active units; White units – inactive.

A. Differentiation

B. Unification

Supported by Grant #7367437 for “Long-term Mobile Monitoring and Analysis of Sleep-Cognition Relationship” from the National Science Foundation's Smart Health and Wellbeing program to M.A.G.

Synaptic deletion during sleep may play a computational role in improving cognitive performance by differentiating and unifying representations

2. Sleep (especially SWS) provides an additional processing stage to the hippocampal representations that were acquired during wake, allowing them to become more parsimonious and consequently boost performance in the subsequent testing phase. This additional stage is based on two processes:

1. Based on our previous NSF-supported modeling (Gluck and Myers, 1993; Moustafa et

Ellenbogen et al., 2006

Plihal & Born, 1997

Lau et al., 2011

Backhaus et al., 2011

Changes in cortical Local Field Potential (LFP) in rats in response to stimulation after a period of wake (Sleep Deprived - SD) compared to sleep (Liu et al., 2010)

Sleep

Gradual learning during wake

Sleep extends pattern differentiation

Sleep sharpens hippocampal input-to-output correlational differences

After sleep-dependent unification and differentiation, each objective is more readily accessed:

Input correlation

Out

put

corr

elat

ion

Contact

==

Itamar Lerner, [email protected] Gluck, [email protected]