Pace-Schott, E. F. & Hobson, J. A. The neurobiology of sleep: genetics, cellular physiology and subcortical networks. Nature Rev. Neurosci. 3, 591–605 (2002).
Hobson, J. A. & McCarley, R. W. The brain as a dream-state generator: an activation–synthesis hypothesis of the dream process. Am. J. Psychiatry 134, 1335–1348 (1977).
Hobson, J. A., Pace-Schott, E. F. & Stickgold, R. Dreaming and the brain: toward a cognitive neuroscience of conscious states. Behav. Brain Sci. 23, 793–842 (2000).
Steriade, M. Active neocortical processes during quiescent sleep. Arch. Ital. Biol. 139, 37–51 (2001).
Borbély, A. A. From slow waves to sleep homeostasis: new perspectives. Arch. Ital. Biol. 139, 53–61 (2001).
Nielsen, T. A. A review of mentation in REM and NREM sleep: 'covert' REM sleep as a possibile reconciliation of two opposing models. Behav. Brain Sci. 23, 851–866 (2000).
Buzsáki, G. The hippocampo–neocortical dialogue. Cereb. Cortex 6, 81–92 (1996).An important and influential theory on state-dependent memory consolidation at the level of the input and output circuits of the hippocampus.
Peigneux, P., Laureys, S., Delbeuck, X. & Maquet, P. Sleeping brain, learning brain. The role of sleep for memory systems. Neuroreport 12, A111–A124 (2001).An important review supporting the theory of sleep-dependent consolidation of neuroplastic changes initiated in waking (see also reference 10).
Stickgold, R. Sleep: off-line memory reprocessing. Trends Cogn. Sci. 2, 484–492 (1998).
Stickgold, R., Hobson, J. A., Fosse, R. & Fosse, M. Sleep, learning and dreams: off-line memory reprocessing. Science 294, 1052–1057 (2001).
Maquet, P. Functional neuroimaging of normal human sleep by positron emission tomography. J. Sleep Res. 9, 207–231 (2000).
Laureys, S. et al. Experience-dependent changes in cerebral functional connectivity during human rapid eye movement sleep. Neuroscience 105, 521–525 (2001).
Maquet, P. The role of sleep in learning and memory. Science 294, 1048–1052 (2001).
Steriade, M. Coherent oscillations and short-term plasticity in corticothalamic networks. Trends Neurosci. 22, 337–345 (1999).
Steriade, M. Corticothalamic resonance, states of vigilance and mentation. Neuroscience 101, 243–276 (2000).
Moruzzi, G. & Magoun, H. W. Brainstem reticular formation and activation of the EEG. Electroencephalogr. Clin. Neurophysiol. 1, 455–473 (1949).
Saper, C. B., Chou, T. C. & Scammell, T. E. The sleep switch: hypothalamic control of sleep and wakefulness. Trends Neurosci. 24, 726–731 (2001).
Rowley, J., Stickgold, R. A. & Hobson. J. A. Eye movement and mental activity at sleep onset. Conscious. Cogn. 7, 67–84 (1998).
Achermann, P. & Borbély, A. A. Low frequency (<1 Hz) oscillations in the human sleep electroencephalogram. Neuroscience 81, 213–222 (1997).A demonstration in the human sleep scalp EEG of the slow oscillation of NREM, an important EEG sign of NREM that was recently identified in depth recordings of the cat by Steriade's group
Simon, N. R., Lopes da Silva, F. H. & Manshanden, I. in Recent Advances in Biomagnetism (ed. Yoshimoto, T.) 373–376 (Tohoku Univ. Press, Tohoku, 1999).
Steriade, M., Nunez, A. & Amzica, F. Intracellular analysis of relations between the slow (<1 Hz) neocortical oscillation and other sleep rhythms of the electroencephalogram. J. Neurosci. 13, 3266–3283 (1993).
Borbély, A. A. & Achermann, P. in Principles and Practice of Sleep Medicine (eds Kryger, M. H., Roth, T. & Dement, W. C.) 337–390 (W. B. Saunders, Philadelphia, Pennsylvania, 2000).
Lisman, J. E. Relating hippocampal circuitry to function: recall of memory sequences by reciptocal dentate–CA3 interactions. Neuron 22, 233–242 (1999).
Rioult-Pedotti, M.-S., Friedman, D. & Donoghue, J. P. Learning-induced LTP in neocortex. Science 290, 533–536 (2000).
Shaffery, J. P., Sinton, C. M., Bissette, G., Roffwarg, H. P. & Marks, G. A. Rapid eye movement sleep deprivation modifies expression of long-term potentiation in visual cortex of immature rats. Neuroscience 110, 431–443 (2002).An important new study showing that the endogenous stimulation of REM sleep might interact with early developmental plasticity of the visual cortex in a manner analogous to external visual input during waking.
Steriade, M. The Intact and Sliced Brain (MIT Press, Cambridge, Massachusetts, 2001).
Borbély, A. A. A two-process model of sleep regulation. Hum. Neurobiol. 1, 195–204 (1982).The initial presentation of a seminal idea in sleep science that is now supported by electrographic, anatomical and molecular findings (see also references 5 and 22).
Czeisler, C. A. & Khalsa, S. S. in Principles and Practice of Sleep Medicine (eds Kryger, M. H., Roth, T. & Dement, W. C.) 353–375 (W. B. Saunders, Philadelphia, Pennsylvania, 2000).
Strecker, R. E. et al. Adenosinergic modulation of basal forebrain and preoptic/anterior hypothalamic neuronal activity in the control of behavioral state. Behav. Brain Res. 115, 183–204 (2000).
Aeschbach, D. et al. Evidence from the waking electroencephalogram that short sleepers live under higher homeostatic sleep pressure than long sleepers. Neuroscience 102, 493–502 (2001).
Werth, E., Achermann P. & Borbély A. A. Fronto-occipital EEG power gradients in human sleep. J. Sleep Res. 6, 102–112 (1997).
Finelli, L. A., Borbély, A. A. & Achermann, P. Functional topography of the human nonREM sleep electroencephalogram. Eur. J. Neurosci. 13, 2282–2290 (2001).An important study showing a frontal predominance of low-frequency EEG rhythms in response to sleep deprivation, which indicates a greater homeostatic need for recovery sleep in the frontal cortex.
Harrison, Y. & Horne, J. The impact of sleep deprivation on decision making: a review. J. Exp. Psychol. Appl. 6, 236–249 (2000).A review on the effects of sleep deprivation on prefrontal functioning.
Beebe, D. W. & Gozal, D. Obstructive sleep apnea and the prefrontal cortex: towards a comprehensive model linking nocturnal upper airway obstruction to daytime cognitive and behavioral deficits. J. Sleep Res. 11, 1–16 (2002).
Balkin, T. J. et al. Bidirectional changes in regional cerebral blood flow across the first 20 minutes of wakefulness. Sleep Res. Online [online] (cited 11 Jul 02) 〈http://www.sro.org/cftemplate/wfsrscongress/indiv.cfm?ID=19998006〉 (1999).
Achermann, P., Werth, E., Dijk, D. J. & Borbély, A. A. Time course of sleep inertia after nighttime and daytime sleep episodes. Arch. Ital. Biol. 134, 109–119 (1995).
Dinges, D. F. in Sleep and Cognition (eds Bootzin, R., Kihlstrom, J. & Schacter, D.) 159–178 (American Psychological Association, Washington DC, 1990).
Llinas, R. & Ribary, U. Coherent 40-Hz oscillation characterizes dream state in humans. Proc. Natl Acad. Sci. USA 90, 2078–2081 (1993).
Anderer, P. et al. Low resolution brain electromagnetic tomography revealed simultaneously active frontal and parietal sleep spindle sources in the human cortex. Neuroscience 103, 581–592 (2001).
Inoue, S., Saha, U. K. & Musha, T. in Rapid Eye Movement Sleep (eds Mallick, B. N. & Inoue, S.) 214–220 (Marcel Dekker, New York, 1999).
Gross, D. W. & Gotman, J. Correlation of high-frequency oscillations with the sleep–wake cycle and cognitive activity in humans. Neuroscience 94, 1005–1018 (1999).
Perez-Garci, E., del Rio-Portilla, Y., Guevara, M. A., Arce, C. & Corsi-Cabrera, M. Paradoxical sleep is characterized by uncoupled gamma activity between frontal and perceptual cortical regions. Sleep 24, 118–126 (2001).
Buchsbaum, M. S., Hazlett, E. A., Wu, J. & Bunney, W. E. Positron emission tomography with deoxyglucose-F18 imaging of sleep. Neuropsychopharmacology 25, S50–S56 (2001).
Maquet, P. Sleep function(s) and cerebral metabolism. Behav. Brain Res. 69, 75–83 (1995).
Maquet, P. et al. Functional neuroanatomy of human slow wave sleep. J. Neurosci. 17, 2807–2812 (1997).
Braun, A. R. et al. Regional cerebral blood flow throughout the sleep–wake cycle. Brain 120, 1173–1197 (1997).The first H215O PET neuroimaging study to include complete pairwise comparisons of waking, NREM sleep and REM sleep (see also references 47, 52 and 53).
Braun, A. R. et al. Dissociated pattern of activity in visual cortices and their projections during human rapid eye-movement sleep. Science 279, 91–95 (1998).An important extension of reference 46 in which evidence is presented that, during REM sleep, internal information is being processed between extrastriate and limbic cortices while they are functionally isolated from the external world in terms of both input (from the striate cortex) and output (through the frontal cortex).
Hofle, N. et al. Regional cerebral blood flow changes as a function of delta and spindle activity during slow wave sleep in humans. J. Neurosci. 17, 4800–4808 (1997).
Andersson, J. et al. Brain networks affected by synchronized sleep visualized by positron emission tomography. J. Cereb. Blood Flow Metab. 18, 701–715 (1998).
Kajimura, N. et al. Activity of midbrain reticular formation and neocortex during the progression of human non-rapid eye movement sleep. J. Neurosci. 19, 10065–10073 (1999).
Nofzinger, E. A. et al. Towards a neurobiology of dysfunctional arousal in depression: the relationship between beta EEG power and regional cerebral glucose metabolism during NREM sleep. Psychiatry Res. 98, 71–91 (2000).
Maquet, P. et al. Functional neuroanatomy of human rapid-eye-movement sleep and dreaming. Nature 383, 163–166 (1996).The first published H215O PET study to compare human REM sleep with other behavioural states, and to show relative activation of limbic and midline subcortical areas and relative deactivation of dorsolateral prefrontal cortex in REM sleep (see also references 46, 47 and 53).
Nofzinger, E. A., Mintun, M. A., Wiseman, M. B., Kupfer, D. J. & Moore, R. Y. Forebrain activation in REM sleep: an FDG PET study. Brain Res. 770, 192–201 (1997).Functional neuroimaging of glucose metabolism in REM sleep compared with waking, in which the anterior paralimbic REM activation area is first specifically identified in sleep (see also references 46, 47 and 52).
Nofzinger, E. A. et al. Changes in forebrain function from waking to REM sleep in depression: preliminary analysis of [18F] FDG PET studies. Psychiatry Res. 91, 59–78 (1999).
Lovblad, K. O. et al. Silent functional magnetic resonance imaging demonstrates focal activation in rapid eye movement sleep. Neurology 53, 2193–2195 (1999).
Nofzinger, E. A. et al. Effects of bupropion SR on anterior paralimbic function during waking and REM sleep in depression: preliminary findings using [18F] FDG PET. Psychiatry Res. 106, 95–111 (2001).
Wu, J. et al. Prediction of antidepressant effects of sleep deprivation by metabolic rates in the ventral anterior cingulate and the medial prefrontal cortex. Am. J. Psychiatry 156, 1149–1158 (1999).
Maquet, P. & Franck, G. REM sleep and the amygdala. Mol. Psychiatry 2, 195–196 (1997).
Peigneux, P. et al. Generation of rapid eye movements during pardoxical sleep in humans. Neuroimage 14, 701–708 (2001).
Fosse, R., Stickgold, R. & Hobson, J. A. Brain–mind states: reciprocal variation in thoughts and hallucinations. Psychol. Sci. 12, 30–36 (2001).The demonstration of a reciprocal relationship between thoughts and hallucinatory activity across five distinct behavioural states in a large longitudinal database of the same subjects.
Hobson, J. A., McCarley, R. W. & Wyzinki, P. W. Sleep cycle oscillation: reciprocal discharge by two brainstem neuronal groups. Science 189, 55–58 (1975).
Ajilore, O. A., Stickgold, R., Rittenhouse. C. & Hobson, J. A. Nightcap: laboratory and home-based evaluation of a portable sleep monitor. Psychophysiology 32, 92–98 (1995).
Stickgold, R., Malia, A., Fosse, R. & Hobson, J. A. Brain–mind states. I. Longitudinal field study of sleep/wake factors influencing mentation report length. Sleep 24, 171–179 (2001).
Fosse, M., Fosse, R., Hobson, J. A. & Stickgold, R. Dreaming and episodic memory: a functional dissociation? J. Cogn. Neurosci. (in the press).
Hasselmo, M. Neuromodulation: acetylcholine and memory consolidation. Trends Cogn. Sci. 3, 351–359 (1999).
Hobson, J. A. Dreaming as Delirium (MIT Press, Cambridge, Massachusetts, 1999).
Solms, M. The Neuropsychology of Dreams: a Clinico-Anatomical Study (Lawrence Erlbaum Associates, Mahwah, New Jersey,1997).
Mesulam, M. M. Principles of Behavioral and Cognitive Neurology (Oxford Univ. Press, Oxford, UK, 2000).
Szymusiak, R. Magnocellular nuclei of the basal forebrain: substrates of sleep and arousal regulation. Sleep 18, 478–500 (1995).
Hobson, J. A., Pace-Schott, E. F. & Stickgold, R. Dream science 2000: a response to commentaries on 'Dreaming and the Brain'. Behav. Brain Sci. 23, 1019–1035 (2000).
Kahn, D., Stickgold, R., Pace-Schott, E. F. & Hobson, J. A. Dreaming and waking consciousness: a character recognition study. J. Sleep Res. 9, 317–325 (2000).
Pace-Schott, E. F. 'Theory of mind,' social cognition and dreaming. Sleep Res. Soc. Bull. 7, 33–36 (2001).
LeDoux, J. E. The Emotional Brain (Simon and Schuster, New York, 1996).
Fosse, R., Stickgold, R. & Hobson, J. A. The mind in REM sleep: reports of emotional experience. Sleep 24, 947–955 (2001).
Merritt, J. M., Stickgold, R., Pace-Schott, E., Williams, J. & Hobson, J. A. Emotion profiles in the dreams of men and women. Conscious. Cogn. 3, 46–60 (1994).
Nielsen, T. A., Deslauriers, D. & Baylor, G. W. Emotions in dream and waking event reports. Dreaming 1, 287–300 (1991).
Paus, T. Primate anterior cingulate cortex: where motor control, drive and cognition interface. Nature Rev. Neurosci. 2, 417–424 (2001).
Damasio, A. R. et al. Subcortical and cortical brain activity during the feeling of self-generated emotions. Nature Neurosci. 3, 1049–1056 (2000).
Liotti, M. et al. Differential limbic-cortical correlates of sadness and anxiety in healthy subjects: implications for affective disorders. Biol. Psychiatry 48, 30–42 (2000).
Pace-Schott, E. F. in Sleep and Dreaming: Scientific Advances and Reconsiderations (eds Pace-Schott, E. F., Solms, M., Blagrove, M. & Harnad, S.) (Cambridge Univ. Press, Cambridge, UK, in the press).
Cahill, L. & McGough, J. L. Mechanisms of emotional arousal and lasting declarative memory. Trends Neurosci. 21, 294–299 (1998).
Porte, H. S. & Hobson, J. A. Physical motion in dreams: one measure of three theories. J. Abnormal. Psychol. 105, 329–335 (1996).
Rye, D. B. Contributions of the pedunculopontine region to normal and altered REM sleep. Sleep 20, 757–788 (1997).
Mori, K., Mitani, H., Fujita, M. & Winters, W. D. Multiple unit activity of dorsal cochlear nucleus and midbrain reticular formation during paradoxical phase of sleep. Electroencephalogr. Clin. Neurophysiol. 33, 104–106 (1972).
Schmahmann, J. D. The role of the cerebellum in affect and psychosis. J. Neurolinguist. 13, 189–214 (2000).
Jouvet, M. The Paradox of Sleep: the Story of Dreaming (MIT Press, Cambridge, Massachusetts, 1999).
Revonsuo, A. The reinterpretation of dreams: an evolutionary hypothesis of the function of dreaming. Behav. Brain Sci. 23, 877–901 (2000).
Haxby, J. V., Hoffman, E. A. & Gobbini, M. I. The distributed human neural system for face perception. Trends Cogn. Sci. 4, 223–233 (2000).
Doricchi, F. & Violani, C. in The Neuropsychology of Sleep and Dreaming (eds Antrobus, J. S. & Bertini, M.) (Lawrence Erlbaum Associates, Mahwah, New Jersey, 1992).
Hobson, J. A. The Dream Drug Store (MIT Press, Cambridge, Massachusetts, 2001).
Siegel, J. The REM sleep–memory consolidation hypothesis. Science 294, 1058–1063 (2001).
Vertes, R. P. & Eastman, K. E. The case against memory consolidation in REM sleep. Behav. Brain Sci. 23, 867–876 (2000).
Roffwarg, H. P., Muzio, J. N. & Dement, W. C. Ontogenetic development of the human sleep–dream cycle. Science 152, 604–619 (1966).
Hobson, J. A. Sleep (Scientific American Library, New York, 1989).
Parmelee, A. H., Wenner, W. H., Akiyama, Y., Schultz, M. & Stern, E. Sleep states in premature and full-term newborn infants. Dev. Med. Child Neurol. 9, 70–77 (1967).
Crick, F. & Mitchison, G. The function of dream sleep. Nature 304, 111–114 (1983).
Marks, G. A., Shaffery, J. P., Oksenberg, A., Speciale, S. G. & Roffwarg, H. P. A functional role for REM sleep in brain maturation. Behav. Brain Res. 69, 1–11 (1995).
Frank, M. G., Issa, N. P. & Stryker, M. P. Sleep enhances plasticity in the developing visual cortex. Neuron 30, 275–287 (2001).
Kirkwood, A. & Bear, M. F. Elementary forms of synaptic plasticity in the visual cortex. Biol. Res. 28, 73–80 (1995).
Cheour, M. et al. Sleep sounds learned by sleeping newborns. Nature 415, 599–600 (2002).
Plihal, W. & Born, J. Effects of early and late nocturnal sleep on declarative and procedural memory. J. Cogn. Neurosci. 9, 534–547 (1997).
Smith, C. Sleep states and memory processes. Behav. Brain Res. 69, 137–145 (1995).
Pavlides, C. & Winson, J. Influences of hippocampal place cell firing in the awake state on the activity of these cells during subsequent sleep episodes. J. Neurosci. 9, 2907–2918 (1989).
Wilson, M. A. & McNaughton, B. L. Reactivation of hippocampal ensemble memories during sleep. Science 265, 676–679 (1994).
Skaggs, W. E. & McNaughton, B. L. Replay of neuronal firing sequences in rat hippocampus during sleep following spatial experience. Science 271, 1870–1873 (1996).
Louie, K. & Wilson, M. A. Temporally structured replay of awake hippocampal ensemble activity during rapid eye movement sleep. Neuron 29, 145–156 (2001).
Poe, G. R., Nitz, D. A., McNaughton, B. L. & Barnes, C. A. Experience dependent phase reversal of hippocampal neuron firing during REM sleep. Brain Res. 855, 176–180 (2000).Shows that rat hippocampal replay of place cells during REM sleep occurs at the peak of theta oscillation — a point that favours LTP — in place cells that correspond to novel environments, but at the trough of theta oscillation — a point that favours long-term depression — for patterns that correspond to familiar environments. The authors suggest that this represents hippocampal consolidation followed by hippocampal erasure as information is transferred, over time, to the neocortex (see also references 103–106 and 108–111).
Nadasdy, Z., Hirase, H., Czurko, A., Csicsvari, J. & Buzsaki, G. Replay and time compression of recurring spike sequences in the hippocampus. J. Neurosci. 19, 9497–9507 (1999).
Hirase, H., Leinekugel, X., Czurko, A., Csicsvari, J. & Buzsaki, G. Firing rates of hippocampal neurons are preserved during subsequent sleep episodes and modified by novel awake experience. Proc. Natl Acad. Sci. USA 98, 9386–9390 (2001).
Siapas, A. G. & Wilson, M. A. Coordinated interactions between hippocampal ripples and cortical spindles during slow wave sleep. Neuron 21, 1123–1128 (1998).
Kudrimoti, H. S., Barnes, G. A. & McNaughton, B. L. Reactivation of hippocampal cell ensembles: effects of behavioral state, experience and EEG dynamics. J. Neurosci. 19, 4090–4101 (1999).
Datta, S. Avoidance task training potentiates phasic pontine-wave density in the rat: a mechanism for sleep-dependent plasticity. J. Neurosci. 20, 8607–8613 (2000).
Sanford, L. D., Silvestri, A. J., Ross, R. J. & Morrison, A. R. Influence of fear conditioning on elicited ponto-geniculo-occipital waves and rapid eye movement sleep. Arch. Ital. Biol. 139, 169–183 (2001).
Dave, A. S. & Margoliash, D. Song replay during sleep and computational rules for sensorimotor vocal learning. Science 290, 812–816 (2000).
Plihal, W. & Born, J. Effects of early and late nocturnal sleep on priming and spatial memory. Psychophysiology 36, 571–582 (1999).
Giuditta, A. et al. The sequential hypothesis of the function of sleep. Behav. Brain Res. 69, 157–166 (1995).
Stickgold, R., Whidbee, D., Schirmer, B., Patel, V. & Hobson, J. A. Visual discrimination task improvement: a multi-step process occurring during sleep. J. Cogn. Neurosci. 12, 246–254 (2000).A study showing a correlation of degree of TDT learning with duration of SWS in the first quarter of the night and REM sleep in the last quarter of the night, indicating a two-step process in the sleep-mediated enhancement of TDT learning (see also references 118–122).
Karni, A. & Sagi, D. Where practice makes perfect in texture discrimination: evidence for primary visual cortex plasticity. Proc. Natl Acad. Sci. USA 88, 4966–4970 (1991).
Stickgold, R., James, L. & Hobson, J. A. Visual discrimination learning requires sleep after training. Nature Neurosci. 3, 1237–1238 (2000).
Karni, A., Tanne, D., Rubenstein, B. S., Askenasy, J. J. M. & Sagi, D. Dependence on REM sleep of overnight improvement of a perceptual skill. Science 265, 679–682 (1994).
Gais, S., Plihal, W., Wagner, U. & Born, J. Early sleep triggers memory for early visual discrimination skills. Nature Neurosci. 3, 1335–1339 (2000).
Mednick, S. et al. The restorative effect of naps on perceptual deterioration. Nature Neurosci. 5, 677–681 (2002).
Walker, M., Brakefield, T., Morgan, A., Hobson, J. A. & Stickgold, R. Practice with sleep makes perfect: sleep-dependent motor learning. Neuron 35, 1–20 (2002).The demonstration of sleep-dependent enhancement of learning on a motor task.
Stickgold, R., Malia, A., Maguire, D., Roddenberry, D. & O'Connor, M. Replaying the game: hypnagogic images in normals and amnesiacs. Science 290, 350–353 (2000).
Stickgold, R., Scott, L., Rittenhouse, C. & Hobson, J. A. Sleep induced changes in associative memory. J. Cogn. Neurosci. 11, 182–193 (1999).
Hartley, D. Observations on Man, His Frame, His Duty and His Expectations (Johnson, London, 1791).
Hobson, J. A., Hoffman, E., Helfand, R. & Kostner, D. Dream bizarreness and the activation–synthesis hypothesis. Hum. Neurobiol. 6, 157–164 (1987).
Maquet, P. et al. Experience-dependent changes in cerebral activation during human REM sleep. Nature Neurosci. 3, 831–836 (2000).The first neuroimaging study to show, in humans, reactivation during sleep of areas corresponding to those activated during a defined preceding waking experience.
Martin, J. H. Neuroanatomy: Text and Atlas 2nd edn (Appleton & Lange, Stamford, Connecticut, 1996).
Hobson, J. A., Stickgold, R. & Pace-Schott, E. F. The neuropsychology of REM sleep dreaming. Neuroreport 9, R1–R14 (1998).
Cantero, J. L., Atienza, M., Stickgold, R. & Hobson, J. A. Nightcap: a reliable system for determining sleep onset latency. Sleep 25, 238–245 (2002).