The Neuroscience of Sleep

Sleep is the natural state of bodily rest, common to all mammals and birds and also seen in many reptiles, amphibians and fish. For most species, regular sleep is essential for survival, yet the specific purposes of sleep are still only partly clear and are the subject of intense research. This volume is comprised of the editors’ selection of the most relevant articles on sleep from the Encyclopedia of Neuroscience, resulting in the first comprehensive collection of introductory articles on the neuroscience of sleep. Articles explore sleep’s impact on neural functioning, sleep disorders, the relation between sleep and other clinical disorders, a look at sleep from a developmental perspective, and more. * Chapters offer impressive scope with topics addressing neural functioning, disorders, development, and more, carefully selected by one of the most preeminent sleep researchers * Richly illustrated in full color with over 100 figures * Contributors represent the most outstanding scholarship in the field, with each chapter providing fully vetted and reliable expert knowledge The Neuroscience of Sleep......Page 2 Copyright Page......Page 5 Table of Contents......Page 6 Contributors......Page 10 Preface......Page 14 INTRODUCTION......Page 16 History of Sleep Research......Page 18 REM Behavior Disorder......Page 21 Further Reading......Page 22 DEFINITIONS AND DESCRIPTIONS OF SLEEP......Page 24 Sleep Architecture......Page 26 Neurophysiological Basis of Sleep Stage Changes......Page 29 Recent Discoveries Regarding the Brain in Sleep......Page 30 Summary......Page 31 Further Reading......Page 32 Confusional Arousals......Page 33 REM Sleep Behavior Disorder......Page 34 Conclusion......Page 35 Further Reading......Page 36 Slow-Wave and Rapid Eye Movement Sleep......Page 37 Three Major Oscillations Define Slow-Wave Sleep......Page 38 Fast Oscillations during REM Sleep and Wakefulness......Page 39 Synaptic Plasticity Induced by Sleep Oscillations......Page 41 Ponto-Geniculo-Occipital Waves......Page 42 Further Reading......Page 43 Non-Rapid Eye Movement Sleep......Page 45 Rapid Eye Movement Sleep......Page 47 Neuroimaging and Dreams......Page 49 Further Reading......Page 50 Introduction......Page 52 The Basics: A Two-Stage Model of Memory Trace Formation......Page 53 Hippocampal Replay of Waking Patterns during Sleep......Page 54 Interactions between Neocortex and Hippocampus during Sleep......Page 55 Conclusions......Page 57 Further Reading......Page 58 Feedback and Rhythms......Page 59 Network Reactivation Increases during Sleep......Page 60 Characterization of Reactivation Properties......Page 62 Reactivation at the Molecular Level......Page 63 Dreams, Waking, and Sleep Mentation......Page 65 Further Reading......Page 66 Hundreds of Genes in the Brain Change Their Expression between Sleep and Waking......Page 67 Three Functional Categories of Waking Genes Are Conserved from Flies to Mammals......Page 69 The Expression of Several Genes Changes after Chronic, but Not Acute, Sleep Loss......Page 71 Conclusions......Page 72 Further Reading......Page 73 ONTOGENY AND PHYLOGENY OF SLEEP......Page 74 Mammalian Sleep......Page 76 Aquatic Mammals......Page 77 Early Ontogeny of Mammalian Sleep......Page 78 Avian Sleep......Page 79 Early Ontogeny of Avian Sleep......Page 80 Corticocortical Connectivity and SWS......Page 81 Comparative Perspectives on the Functions of Sleep......Page 82 Sleep in Invertebrates......Page 83 Further Reading......Page 84 Adolescent Development and the Waking Electroencephalogram......Page 85 Adolescent Development and the Sleeping EEG......Page 86 Homeostatic Process (Process S)......Page 87 Circadian Timing (Process C)......Page 88 Interaction of Process S and Process C......Page 89 Sleep Habits and Adolescent Development......Page 90 Further Reading......Page 92 Treatment of Circadian Rhythm Disturbances......Page 93 Medical and Psychiatric Illness and Insomnia......Page 94 Pharmacological therapy......Page 95 Significance of SDB in development......Page 97 Conclusion......Page 98 Relevant Websites......Page 99 CONTROL OF SLEEP AND SLEEP STATES......Page 100 Thalamic Pacemakers for Sleep Spindle Oscillations......Page 102 Dialog between the Thalamus and Cortex: Emergence of Large-Scale Synchrony......Page 104 Further Reading......Page 105 Electrocortical and Behavioral Arousal: The Ascending Reticular Activating System......Page 106 New Perspective on the ARAS......Page 107 Sleep - Turning the Ascending Arousal System Off......Page 108 Flip-Flop Switch Stability and the Lateral Hypothalamic Area......Page 109 Circadian Regulation of Sleep......Page 110 Hypothalamic Circadian Integrator for Sleep-Wake Regulation......Page 111 Further Reading......Page 112 Physiology of Histaminergic Neurons......Page 114 Amines......Page 115 Purines......Page 116 H1 Receptors......Page 117 H2 Receptors......Page 118 H3 Receptors......Page 119 Histamine and Synaptic Plasticity......Page 120 Control of Waking and Arousal......Page 121 Conclusion......Page 122 Further Reading......Page 123 TNF-alpha in Sleep Regulation......Page 124 TNF Cell Biology......Page 126 Brain Organization of Sleep and TNF-alpha......Page 127 Regulation of REMS by Prolactin and Related Molecules......Page 128 A Proposed Mechanism for PRL-Enhanced REMS......Page 129 Further Reading......Page 130 Projections and Role of Noradrenergic Locus Coeruleus Neurons in Arousal......Page 131 Projections and Role of Serotonergic Raphe Neurons in Waking and Sleeping......Page 134 Further Reading......Page 135 Cholinergic Systems in the Brain......Page 136 Cholinergic Systems and Cortical Activation of Waking......Page 137 Cholinergic Systems and REM Sleep......Page 138 Further Reading......Page 139 Sleep Regulation: Wake-Active Dopamine Neurons......Page 140 Dopaminergic Disorders and Sleep: PD......Page 142 Dopamine and Stimulant-Induced Arousal......Page 143 Further Reading......Page 144 SLEEP AND AROUSAL STATES......Page 146 Wiring Diagram......Page 148 Intrinsic Properties......Page 149 Slow-Wave Sleep......Page 150 Muscle Tone and Locomotion......Page 151 Stimulants......Page 152 Arousal and Sleep Disorders......Page 153 Further Reading......Page 154 Etiology......Page 155 Nightmare Frequency versus Nightmare Distress......Page 156 Nightmares and Sleep Physiology......Page 157 Case Report - Adult......Page 158 Conclusion and Future Directions......Page 159 Further Reading......Page 160 Brain Death......Page 161 Coma......Page 162 MCS......Page 163 Coma......Page 164 Brain Death......Page 165 VS......Page 166 LIS......Page 169 Further Reading......Page 170 CIRCADIAN......Page 172 Hierarchy and Cell-Autonomous Nature of Circadian Clocks in Mammals......Page 174 Functional Differences between Circadian Clocks in the Suprachiasmatic Nucleus, Peripheral Tissues, and Fibroblasts......Page 175 The Search for Critical Output Signals of the Suprachiasmatic Nucleus......Page 177 Further Reading......Page 179 Three Stages of Sleep Development......Page 180 Sleep Regulation......Page 181 Development of the Circadian System......Page 182 Sleep and Circadian Development Continues throughout Life......Page 183 Further Reading......Page 184 Hundreds of Genes in the Brain Change Their Expression between Sleep and Waking......Page 185 Three Functional Categories of Waking Genes Are Conserved from Flies to Mammals......Page 187 The Expression of Several Genes Changes after Chronic, but Not Acute Sleep Loss......Page 189 Conclusions......Page 190 Further Reading......Page 191 Circadian Rhythm Sleep Disorders......Page 192 Diagnosis......Page 193 Pathophysiology......Page 194 Summary......Page 196 Further Reading......Page 197 Subjective Measures of Fatigue and Alertness......Page 198 Electroencephalographic and Ocular Measures......Page 199 Interindividual and Intraindividual Variability......Page 200 Forced Desynchrony......Page 201 Accident Risk......Page 202 Further Reading......Page 203 PHARMACOLOGY OF SLEEP......Page 204 Modafinil Pharmacology......Page 206 Amphetamine Pharmacology......Page 207 Caffeine......Page 208 Further Reading......Page 210 General Pharmacology of Sleep......Page 212 Adenosine as a Potential Target of Hypnotics......Page 214 Adenosine Receptors as Therapeutic Targets......Page 216 Nucleoside Transporters as Therapeutic Targets......Page 217 Allosteric Modulators as Therapeutic Agents......Page 218 Allosteric Modulators of the A3 Adenosine Receptor Subtype......Page 219 Adenosine as a Therapeutic Agent for Other Diseases......Page 220 Further Reading......Page 221 SLEEP FUNCTION......Page 222 Key Hormones of the Anterior Pituitary......Page 224 Thyroid-Stimulating Hormone......Page 226 Further Reading......Page 227 Human Models......Page 228 Animal Models......Page 229 Sleep Deprivation in Humans......Page 230 Further Reading......Page 232 Thermoregulatory Responses to Thermal Challenge Are Influenced by Sleep......Page 233 Temperature-Sensing Neurons in the Hypothalamus Control Body Temperature......Page 234 Acute Sleep Deprivation: Interactions with Circadian Control of Body Temperature......Page 235 Thermoregulatory Changes Influence Sleep Onset and Sleep Maintenance......Page 236 Further Reading......Page 237 Dreaming......Page 238 Cardiovascular Reflexes Mediating Autonomic Activity......Page 239 Measures of Autonomic Cardiovascular Regulation......Page 240 REM Sleep and Heart Rate/Rhythm......Page 241 REM Sleep and Autonomics in the Setting of Cardiovascular Disease......Page 242 Further Reading......Page 243 Memory Categories......Page 245 Memory Stages......Page 246 Sleep and Memory Encoding......Page 247 Declarative Memory......Page 248 Motor learning......Page 250 Procedural memory and daytime naps......Page 251 Overnight Reorganization of Memory Representations......Page 253 Further Reading......Page 255 History of Sleep Research......Page 256 Sleep Deprivation and the Propensity to Fall Asleep......Page 257 Sleep Deprivation and Basic Cognitive and Psychomotor Performance......Page 258 Sleep Deprivation, Mood, and Emotion......Page 259 Polysomnography and Waking EEG and Electrooculography Effects of Chronic Sleep Restriction......Page 260 Differential Neurobehavioral Vulnerability to Sleep Loss......Page 262 Further Reading......Page 263 Functional Neuroimaging Studies of Sleep Deprivation......Page 264 Learning Tasks......Page 265 Working Memory......Page 266 Compensatory Recruitment Hypothesis......Page 267 Further Reading......Page 268 Changes in Napping Behaviors across the Life Span......Page 269 Physiological Evidence for 'Nap Zone'......Page 270 Behavioral Markers for the 'Nap' Zone......Page 271 Napping in Sleep-Deprived Conditions......Page 272 Summary......Page 273 Relevant Website......Page 274 SLEEP DISORDERS......Page 276 Sleep Apneas Are Neurally Mediated......Page 278 Obstructive Sleep Apnea......Page 279 Obstructive Sleep Apnea......Page 280 Central Sleep Apnea......Page 281 Obstructive Sleep Apnea......Page 282 Future Directions......Page 283 Further Reading......Page 284 Dissociated REM Sleep in Narcolepsy......Page 285 Neurobiology of the Hypocretin Systems and Relevance to Narcolepsy......Page 286 Is Narcolepsy an Autoimmune Disorder?......Page 289 Narcolepsy without Cataplexy......Page 290 Therapies for Narcolepsy......Page 291 Further Reading......Page 292 Clinical Features of Cataplexy......Page 293 Physiological and Pathophysiological Aspects of Cataplexy......Page 295 Cataplexy and Hypocretin/Orexin Deficiency......Page 296 Relevant Website......Page 299 Historical Notes and Current Prevalence......Page 300 Stages of Sleeping Sickness, Treatment, and Neurological Signs......Page 301 Neuropathological Findings......Page 302 Neuroinflammation in Trypanosoma brucei-Infected Rodents......Page 303 Involvement of Neural Centers That Regulate Circadian Rhythms and Sleep......Page 304 The Blood-Brain Barrier and Implications for Treatment......Page 305 Further Reading......Page 306 DREAMING......Page 308 Carryover Effects of REM Dreaming......Page 310 Sleep paralysis episodes......Page 311 Effects of REM deprivation on consolidation......Page 312 Dream Content and Memory Consolidation......Page 313 REM deprivation and depression......Page 314 Conclusion......Page 315 Further Reading......Page 316 Changes in the Brain and in the Body during Sleep; Changes in Consciousness......Page 317 Formal Analysis of Dream Content......Page 318 Input-Output Gating......Page 320 Efferent Copy Generation......Page 321 Convergence of Physiology, Psychology, and Philosophy......Page 322 AIM: An Integrated Model of the Human Brain Mind......Page 323 Further Reading......Page 324 The Activation-Synthesis Model......Page 325 Hallucinatory Quality......Page 326 Personal Motives, Values, and Emotions......Page 327 Circadian Influences on Sleep Mentation......Page 328 Sleep State Ambiguity and Mentation Features......Page 329 Further Reading......Page 330 Dream Recall and Nightmare-Related PTSD......Page 331 Victims of War and Violence......Page 332 Sensitization Hypothesis of PTSD Nightmares......Page 333 Potential Psychophysiological and Neurobiological Underpinnings of PTSD-Related Nightmares......Page 334 Imagery Rescripting and Rehearsal......Page 335 Other Pharmacological Agents......Page 336 Relevant Websites......Page 337 Nightmare Prevalence and Correlates......Page 338 Boundary permeability......Page 339 Threat simulation......Page 340 REM sleep desomatization......Page 341 Affective network dysfunction......Page 342 Further Reading......Page 344 Manipulation of Presleep Experience in the Sleep Laboratory......Page 345 Incorporation of Presleep Experience into Sleep Onset Dreaming......Page 346 Form of Waking Experience Incorporations......Page 347 Emotional Value and Waking Experience Incorporation......Page 348 Recent Memories, Dreaming, and Memory Consolidation......Page 349 Neural Basis for Dream Incorporation of Recent Events: Connection to Declarative Memory Consolidation?......Page 350 Further Reading......Page 351 Subject Index......Page 352 Section I: Introduction. History of sleep research / R. Pelayo and C. Guilleminault Section II: Definitions and descriptions of sleep. Sleep architecture / E. F. Pace-Schott ; Behavior and parasomnias (RSBD) / M. W. Mahowald, M. A. Cramer-Bornemann, and C. H. Schenck ; Sleep oscillations and PGO waves / M. Steriade ; PET activation patterns / T. T. Dang-Vu, M. Desseilles, P. Peigneux, S. Laureys, and P. Maquet ; Hippocampal-neocortical dialog / J. J. Chrobak, A. Sirota, and G. Buzáski ; Network reactivation / S. Ribeiro and M. Nicolelis ; Gene expression / C. Cirelli and G. Tononi Section III: Ontogeny and phylogeny of sleep. Phylogeny and ontogeny of sleep / J. A. Lesku, D. Martinez-Gonzalez, and N. C. Rattenborg ; Sleep in adolescents / I. Tarokh and M. A. Carskadon ; Sleep in aging / M. Rissling and S. Ancoli-Israel Section IV: Control of sleep and sleep states. Thalamic regulation of sleep / A. Destexhe and T. J. Sejnowski ; Hypothalamic regulation of sleep / P. M. Fuller and J. Lu ; Histamine / H. L. Haas, O. Selbach, and O. A. Sergeeva ; Cytokines and other neuromodulators / J. M. Krueger, L. Churchill, and D. M. Rector ; Locus coeruleus and Raphe nucleus / B. E. Jones ; Acetylcholine / B. E. Jones ; Dopamine / P. M. Fuller and J. Lu Section V: Sleep and arousal states. Reticular activating system / E. Garcia-Rill ; Nightmares / M. Schredl ; Coma / S. Laureys, M. Boly, G. Moonen, and P. Maquet Section VI: Circadian. Circadian regulation by the suprachiasmatic nucleus / D. J. Earnest ; Sleep: development and circadian control / H. C. Heller and M. G. Frank ; Sleep and waking in drosophila / C. Cirelli and G. Tononi ; Genetics of circadian disorders in humans / A.-M. Chang and P. C. Zee ; Circadian rhythms in sleepiness, alertness, and performance / J. D. Minkel and D. F. Dinges Section VII: Pharmacology of sleep. Modafini, amphetamines, and caffeine / C. Ballas and D. F. Dinges ; Adenosine / R. Basheer, J. T. McKenna, and R. W. McCarley Section VIII: Sleep function. Endocrine function during sleep and sleep deprivation / J. M. Mullington ; Immune function during sleep and sleep deprivation / J. M. Mullington ; Thermoregulation during sleep and sleep deprivation / R. Szymusiak ; Autonomic dysregulation during REM sleep / S. M. Caples and V. K. Somers ; Sleep-dependent memory processing / M. P. Walker ; Behavioral change with sleep deprivation / J. D. Minkel, S. Banks, and D. F. Dinges ; Sleep deprivation and brain function / S. P. A. Drummond and B. S. McKenna ; Napping / S. C. Medrick and S. P. A. Drummond Section IX: Sleep disorders. Sleep apnea / S. C. Veasey ; Narcolepsy / E. Mignot and L. Lin ; Cataplexy / S. Nishino ; Sleeping sickness / M. Bentivoglio and K. Kristensson Section X: Dreaming. Theories of dream function / D. Kuiken ; Sleep and consciousness / J. A. Hobson ; REM/REM differences in dream content / J. S. Antrobus and E. J. Wamsley ; Dreams and nightmares in PTSD / A. Germain and A. Zadra ; Theories and correlates of nightmares / T. Nielsen and R. Levin ; Incorporation of waking events into dreams / E. J. Wamsley and R. Stickgold.