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• Psychology of (Psych 133) – Prof. Allison Harvey Sleep and • Sleep and Psychological Disorders Laboratory – Prof. Matt Walker • Sleep and Neuroimaging Laboratory Fall 2014 • Psychology of Dreams (Psych 106) – Prof. Eleanor Rosch

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Why Sleep is Interesting Diagnosis of Sleep • Puzzle of Function • Role in Learning, Memory • Overt Behavior • , Other Sleep Disorders – Relaxation • Lapse in Consciousness – Prone Posture – Consciousness as Wakefulness – Slow, Even Breathing – Contrast with “Dreamless Sleep” • Subjective Experience • Conscious of Dreams? – Interruption of Stream of Consciousness • Unconscious Processing – Disorientation upon Awakening • Mind-Body Problem – Memory Failure – Physiological Correlates of Sleep/Dreams – Recall 3 4

The EEG Spectrum Hugdahl (1995)

Polysomnography Rechtschaffen & Kales (1968)

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1 Prominent EEG Bands The Waking EEG

• EEG Bands • Alpha Activity –Slow –Delta(0.5-4 Hz) – High-Amplitude – Theta (5-7 Hz) –Rhythmic – Arousal – Alpha (8-12 Hz) • Inverted-U –Beta(18-30 Hz) • Beta Activity –Fast

– Gamma (30-50 Hz) Wikipedia Hugo Gamboa Dez 2005 – Low-Amplitude – High Gamma (>250 Hz) – Desynchronized – “Looking”

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Drowsiness Descending Stage 1 • EEG – Alpha Disappears • Eyes Closed – Desynchronized Activity • Low Body Motility • EMG Muscle Relaxation • Decrease in Body • EOG Slow Rolling Eye Movements Temperature • Lack of Behavioral • EEG Alpha Activity Response – Initial Increase • Denial of Sleep on – Subsequent Decrease Awakening?

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Stage 2 Stage 3

• EMG Relaxation • EMG Relaxation • EOG SREMs • EOG SREMs • EEG Changes • EEG Delta Activity – Spindles • Awakening to Loud – K-complexes Noise, Name

11 12 K-Complex Spindle

2 Stage 4 Stage REM • EEG Resembles Stage 1 • EMG Relaxation • EMG Relaxation • EOG SREMs – • EEG Delta Activity – • Awaken Groggy • EOG REMs () – Conjugate • ANS Arousal – High EKG

13 • Awaken Alert 14

Stage NREM (SWS) Stages of Sleep Sleep Architecture “Descending” Stage 1 Stages 2, 3, 4 in Disorders of Consciousness • EMG Relaxed Cologan et al. (2009) • EOG SREMs Coma Vegetative State • EEG Slowing • EEG Slowing • Sleep Stages • “Slow-Wave Sleep” – Delta, Theta – Spindles, K-complexes – Stages 3, 4 • “Spindle Coma – Alternating REM, SWS – Better Prognosis? Stage REM “Ascending” Stage 1 • EMG Relaxed • EOG REMs • EEG Fast 15 16

Nervous System Activity in Sleep

SWS REM Autonomic Heart Rate Slow Decline Variable Respiration Slow Decline Variable Thermoregulation Maintained Impaired Brain Temperature Decreased Increased Cerebral Blood Flow Reduced High Somatic Postural Tension Slow Decline Eliminated Patellar Reflex Normal Suppressed Phasic Twitches Reduced Increased Eye Movements Infrequent, Slow Rapid, Conjugate 17 Hobson (1995) 18

3 Succession of Sleep Stages Dement (1976) Consciousness During Sleep

• Aspects of Information-Processing – Pickup Information from Environment – Integrate with Pre-Existing Knowledge – Respond Meaningfully while Asleep – Remember in Morning • Evidence (Largely Anecdotal) – Body Movements – Awakening to Meaningful Sounds – Awakening at Preselected Times 19 – Dream-Incorporation 20

Sleep Stage and Reports of Sleeping Falling Asleep Webb (1975) Wyatt et al. (1994) • Paired Associates (hot-cold) 100 – Lights out, 1/minute 90 80 – Repeat word pair 70 60 Don't Know 50 Awake • Repetition failure 40 Asleep 30 – 15 secs of alpha-free Stage 1

% of Subjects of % 20 10 – further 30 secs or 10 min 0 Awake 1 2 3, 4 • Awakening Sleep Stage – Recall and Recognition Tests – 10 Cycles 21 22

Retrograde Amnesia in Sleep Wyatt et al. (1994) Explicit vs. Implicit Memory

• Explicit Test – Associate-Cued Recall • Implicit Test – Free Association

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4 Explicit vs. Implicit Memory: Dreaming and REM Sleep Last 2 Minutes Before Sleep Aserinsky & Kleitman (1953) Wyatt et al. (1994) • Observed Cyclical Shift from SREM to REM 90 80 – 10-20 Minute Bursts 70 60 30sW – c. Every 90 Minutes Baseline 50 10mW 40 30sS • Awaken Subjects in Various Sleep Stages 30 10mS 20 • “Were You Dreaming?”

Corrected % Targets Corrected % 10 – Awakened in REM, 74% 0 Explicit Implicit – Awakened in NREM, 17% Memory Test • Sleep Not a “Passive” Phenomenon 25 – 2 Kinds of Sleep 26

Sleep Stage and Dream Recall Mental and Behavioral Dement (1976) Activity During Sleep

• Descending Stage 1 100 – Hypnagogic Reverie 80 • Stage NREM 60 REM – Reverie, Thoughts, Images 40 NREM – , Sleep Talking, Night Terrors 20 % DreamReports • Stage REM 0 A&K (1953) 9 Studies – Dreams, Study • Ascending Stage 1 27 – Reverie 28

Hypnagogic Reverie Night Terrors Schacter (1976) (Pavor Nocturnus, Incubus) Gastaut & Broughton (1964) • Imagery in Descending Stage 1 – Visual • Behavioral • Patterns, Static Objects, Complex Scenes – Inarticulate Scream, Cry for Help, Fear – Auditory – Unable to account for distress • Images, Music, Own Name – Return to (peaceful) sleep • “Myoclonic Jerk” – No memory upon awakening – Accompanying imagery • Physiological – Muscles-Tendons Antagonism – Intense ANS Discharge • Hypnopompic Reverie (Ascending Stage 1) – Arousal from Early Stages 3, 4 – Sleep Paralysis 29 • Not 30

5 Nightmares Nightmares and Night Terrors Hartmann (1984) Hartmann (1984)

• Frightening Dream – Awakens Sleeper • Stage REM – Late in Night

Fuseli, the Nightmare (1781)

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Sleepwalking (Somnambulism) Kales et al. (1966) Sleeptalking (Somniloquy) Arkin et al. (1978)

• Speech, Other Meaningful Sounds • Stage NREM (SWS) – No Awareness on Awakening • Random, Purposelessness • Children vs. Adults • Dexterity • REM, Concordant with Dream Contents • Episodic – Syntax, Semantics • Affect – Monologue, One-Sided Conversation • Development • NREM, Discordant with Dream Contents – Children – Aphasic? – Adults 33 34

Parasomnias Early Studies of Sleep-Learning

In REM In NREM • Thorndike (1916), Morse Code • REM Sleep Behavior • Sleepwalking – Learning Disorder • Night Terrors – Fatigue • Recurrent Isolated • – Surreptitious Practice Sleep Paralysis • Restless Leg Syndrome • Fox & Robins (1952) – Chinese-English Paired-Associates • Correct Pairs, Incorrect Pairs • Music as Control

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6 Sleep and Waking Learning Analysis of Sleep Learning Fox & Robins (1952) Simon & Emmons (1955)

100 12 90 80 10 70 Hear 8 60 50 Recall 6 40 Recog 30 4 % of Items 20 10 2 0 Trials to Learn AM in Trials 0 Correct Music Incorrect Pair Type Sleep Stage

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Alpha Activity in Memory Simon & Emmons (1956)

35 “Sleep learning is possible, 30 25 to the extent that the subject 20 remains awake” 15 Simon & Emmons (1955) % of Items% of 10 5 0 10Sec None No Hear Alpha Activity Following Item

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Explicit and Implicit Memory Homophone Priming for Sleep Learning Wood et al. (1992) Wood et al. (1992)

• Night 1, Homophone Pairs 80 – War-Peace, Tortoise-Hare 70 60 – Homophone Spelling 50 Targets 40 • Night 2, Category-Instance Pairs Lures 30 – Metal-Gold, Animal-Horse 20 10 Standardized Scores Standardized – Category Instance Generation 0 • Signs of Arousal Sleeping Waking Condition – Body Movement – EEG Alpha, SREMs 41 42

7 Category-Instance Priming Wood et al. (1992)

120 “Sleep learning is possible, 100 to the extent that the subject 80 Targets 60 remains awake” Lures 40 Simon & Emmons (1955) 20 Standardized Scores Standardized 0 Sleeping Waking Condition

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Memory for Dreams Factors Affecting “Consolidation”

• Problem • Low Arousal During Sleep – High Frequency of Dreams in Stage REM • REM Awakenings – 4-5 Epochs of REM in Night’s Sleep – Abrupt vs. Gradual Arousal – Why Remember Only 1 Dream (At Most)? – Distraction • Dream Occurs in Primary/Working Memory • Salience – REM Awakening Permits Immediate Readout – Consolidation – Retrieval 45 46

Sleep Stage and Mentation Sleep Stage and Dream Recall Foulkes (1962); Foulkes & Schmidt (1983) Aserinsky & Kleitman (1953); Dement (1976)

100 100

80 80

60 REM 60 REM 40 NREM 40 NREM

20 20 % of Awakenings of % % DreamReports

0 0 A&K (1953) 9 Studies 1962 1983 Study Study

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8 Normative Characteristics of Dreams Snyder (1970) Characteristics of NREM Mentation • Vivid, Complex Imagery • Stage 1 (NREM) • Temporal Progression – Reverie, Daydreaming • Primarily Visual • Stage 2 • Familiar Setting – Imageless Thoughts, Reverie • Dreamer as Central Character • Stage 3 • Moderate to High Credibility – Imageless Thoughts, Floating Images • Fairly Coherent • Stage 4 • Flat Affect – Incoherent Reports (Sleep Inertia?)

– Nightmares 49 50

Analysis of Sleep Mentation Dream Content Foulkes & Schmitt (1983) Hall & Van de Castle (1966) • Dreams Constant Despite Cultural Change

Quality REM NREM • No Changes in Content Across Adulthood • Stable Patterns of Differences Across Mental 93% 67% Cultures Content % "Dreams" 80% 40% – Characters: Women, M=F; Men, M>>F – Aggression>Friendliness Dreams 74% 27% – Misfortune>Good Fortune Length 5.5 1.3 – Emotion: Negative>Positive • Individual Differences Consistent with 51 Waking Personality 52

The Interpretation of Dreams Dreams as Wish Fulfillments Freud (1900) Freud (1900)

“I shall demonstrate that there is a • The Dream of Irma psychological technique which makes it – Injection of Trimethylamine possible to interpret dreams, and that • Anna’s Dream (a Paraphrase) on the application of this technique, – Stwabewwies! every dream will reveal itself as a psychological structure, full of significance….”

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9 Vocabulary of the Dream Theory Common Dream Symbols Freud (1900) Freud (1900), Chapter 6

• Day Residues • A Hat as a Symbol of a Man (or of Male Genitals) • Manifest vs. Latent Content • A “Little One” as the Genital Organ – “A Little One Being Run Over” as a Symbol of Sexual Intercourse • • The Genitals Represented by Buildings, Stairs and Shafts – Displacement • The Male Organ Represented by Persons – Condensation • The Female Organ by a Landscape – Visual Representation • Dreams of Castration in Children • Urinary Symbolism – Secondary Revision

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Topographical Theory of the Mind Dreaming Begins with “PGO Waves” Freud (1900)’s “Compound Instrument” Brooks & Bizzi (1963), after Jouvet, Michel, & Courjon (1959)

• Implanted Microelectrodes – Cats, Rodents • Activity During REM Sleep – Not during SWS, Waking • Perceptual, Mnemic Systems • Sequential Activation • Motor Activity 1. Pons • Systems Ucs, Pcs, Cs 2. Lateral Geniculate Nucleus Types of 3. Occipital Cortex Executive • Cathexis and Anticathexis Trigger, Transfer Modulatory • Replaced by Functional Theory (1923) Aminergic (NE, 5-HT) (Ach) – Id, Ego, Superego 57 58

AIM Model of Consciousness Activation-Synthesis Theory of Dreams “An Integrative Theory of Mind-Brain States” Hobson & McCarley (1977) Hobson et al. (1990, 1992, 1995, 1999, 2000, 2001, 2014)

• Cyclic Activation of Cortex Psychological Level Physiological Level – Controlled by Biological Clock • Alertness • Cortical Activation – Consequences of Activation –RAS – Neural Firing Rate in Brainstem • Feedback of Eye Movements • Motor Commands • Input Source • Sensory/Motor Channels • Vestibular Activity – Internal vs. External – Input-Output Gating • ANS Activity • Information Processing • Neuromodulatory Balance – Aminergic • Automatic Synthesis of Imagery – Attentiveness – Corresponding to Sensory Activity • (NE) • (5-HT)

• Dreams are Essentially Meaningless 59 – Cholinergic Neurons 60 • (Ach)

10 Brainstem Neuromodulatory Systems Noradrenergic Neuromodulatory System Hobson (2001) Hobson (2001)

• Locus Coeruleus • Acetylcholine – Spinal Cord • Noradrenergic –Brainstem – Cerebellum • Serotonergic – • (Dopaminergic) – Not Visible – Subthalamus – Not Yet in AIM Model – Limbic System – Neocortex

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Seratonergic Neuromodulatory System Acetylcholine Neuromodulatory System Hobson (2001) Hobson (2001)

• Raphe Nuclei • Sources – Spinal Cord – Pons –Brainstem • Thalamus – Cerebellum • Subthalamus • Basal Forebrain – Thalamus • Limbic System – Subthalamus – Basal Forebrain – Limbic System • Neocortex – Neocortex • Limbic System

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Dopaminergic Neuromodulatory System The AIM State-Space Hobson (2001) Hobson (1990)

LowActivation High • Midbrain – Limbic System

– Neocortex High – Extrapyramidal Aminergic Motor System • Not Involved in Modulation Sleep and Dreams

• Relevant to External Psychedelic States Input Source –LSD High 65 Cholinergic Internal 66

11 Waking Consciousness in the AIM State-Space NREM in the AIM State-Space

LowActivation High LowActivation High

Waking Waking

High High Aminergic Aminergic

NREM

Modulation Modulation

External External

Input Source Input Source High High Cholinergic Internal 67 Cholinergic Internal 68

REM in the AIM State-Space in the AIM State-Space

LowActivation High LowActivation High

Waking Waking

High High Aminergic Aminergic

NREM NREM

Modulation Modulation

External External REM REM Input Source Input Source High High Cholinergic Internal 69 Cholinergic Internal 70

AIM and Sleep Consequences of REM Modulation Hobson (1990) •Waking – High Activation, External Information • High Levels of Cortical Activation – Aminergic > Cholinergic – Lots of Mental Activity • NREM Sleep • Shift from External to Internal Inputs – Low Activation, Internal Information – Mental Activity Dominated by Memory – Aminergic = Cholinergic – No Behavioral Outputs • REM Sleep • Low Aminergic Activity – High Activation, Internal Information – Poor Memory on Waking – Cholinergic > Aminergic

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12 The AIM Model Beyond Sleep and Dreams Implications of AIM Model Hobson et al. (2000) • Brainstem Critical for Dreams • Activation – Reticular Activating System – Normal Consciousness (High) • Dreams Meaningless – Quiet Waking (Low) – Contents a Product of Random Activation • Input • Focus on Formal Properties of Thought – Daydreams, Fantasies – Similar to Waking Thought • Modulation – Psychedelic Drugs But Are Dreams Really Devoid of Meaning? – Depression 73 74

Brain Damage and Dreaming Are Dreams and REM Dissociable? Solms (1997) Solms (2000) • Dreams Occur in NREM as well as REM • 200/332 Patients with No Changes – 5-10% of NREM Awakenings  Dream Report – Dosolateral Prefrontal Cortex • Indistinguishable from REM Dream Reports – Sensorimotor Cortex – 5-30% of REM Awakenings  No Dream – Primary Visual Cortex • Forebrain Mechanisms Critical for Dreaming • 121/132 Patients Lost All Dreaming – Cholinergic Systems Control REM – Parietal Lobes (Spatial Representation) – Frontal/Dopaminergic Systems Control Dreaming – Frontal-Limbic Region (Executive Functions) • A Double Dissociation? • 2 Patients Lost Visual Imagery – Pontine Damage Suppresses REM, not Dreams – Damage in Visual Association Cortex 75 – Frontal Damage Suppresses Dreams, not REM76

A New Dream Theory A “Dream-On” Mechanism? Solms (2000) Solms (2000)

• Evidence for a Dopaminergic Hypothesis • Mesocortical/Mesolimbic – Forebrain Transection Eliminates Dreaming System • Interrupts Mesocortical/Mesolimbic Dopamine System – Vental Tegmental Area (VTA) • No Effect on REM Sleep – Amygdala – L-dopa Stimulates Vivid Dreams, Nightmares – Prefrontal Cortex – Haldol Inhibits Frequent/Vivid Dreams • Dreams as Wish-Fulfillments? • Dream-Generation Process – Goal-Directed Behavior • A “Seeking System” – Cerebral Activation During Sleep – Reward/Pleasure thebrain.mcgill.ca • Many Different Origins, Not Just Pontine Activity • Neuro-Psychoanalysis

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13 Hobson and Solms Compared and Contrasted A Cognitive Theory of Dreams Foulkes (1985) Domhoff (2005) • Differences – Role of PGO Waves • Development of Dreaming – Association of Dreaming with REM – Dream Recall Minimal Before Age 9 – Role of – Low Levels of Negative Content in Children • Cholinergic/Adrenergic vs. Dopamine • Dreaming Instigation • Similarities – Random Activation of Memory Structures – Dreaming as Psychosis • Episodic, Semantic – Dream Content Insignificant • Dream-Production system – Need to Polarize Debate – Organizes Random Elements 79 • Coherent Dream Experience 80

Sources of Mnemonic Activation in Sleep Dream-Production System Foulkes (1985) Foulkes (1985) • Dreaming as Thinking • External or Internal Stimuli – Visual-Spatial Thinking Skills – Spontaneous Brain Activity • Right Hemisphere? • Associations with Activated Memories – Syntactical Rules, Script Knowledge • Left Hemisphere? – Looser in Sleep • One Dream-Production at All Stages • Voluntary Retrieval – Cortical Excitation Greater in REM – Absent • Developmental Trends (“Piagetian”) – No Dreams During Sensory-Motor Period 81 • Amnesia as Encoding Failure 82

The Meaning of Dreams Dream Content Revisited Foulkes (1985) Domhoff (2001, 2005) • Uncertain Relationship b/ Dreams, Physiology • Indicative Meaning – REM and Dreaming (REM Deprivation) – Reflect Mind of Dreamer – Unusual Eye Movements and Bizarreness • Random Sampling • Faithfulness to Everyday Waking Life • Personality – Commonplace, Familiar Settings • Life Situation – Low Degree of Drama – Low Degree of Bizarrenes – Little Emotion – Thinking Generally Coherent 83 • Laboratory vs. Home Environments 84

14 Neurocognitive Theory of Dreams Domhoff (2001, 2005)

• Neural Network for Dreaming – Forebrain • Limbic System • Inferior Parietal Cortex • Dreaming as Cognitive Achievement – Develops Over First 9 Years of Life • Dreams Continuous with Waking Life – Continuity Principle

– Repetition Principle 85 86

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