Review
doi: 10.1016/B978-0-444-52006-7.00009-5.Neurobiology of waking and sleeping
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- PMID:21056184
- PMCID: PMC5548106
- DOI: 10.1016/B978-0-444-52006-7.00009-5
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Review
Neurobiology of waking and sleeping
Barbara E Jones. Handb Clin Neurol.2011.
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Sleep–wake state substrates. Sagittal schematic view of the human brain depicting neurons with their chemical neurotransmitters and pathways by which they influence cortical activity or behavior across the sleep–wake cycle. Neurons which are active during waking (red symbols) include cells with ascending projections toward the cortex, which stimulate cortical activation, and cells with descending projections toward the spinal cord, which stimulate behavioral arousal with postural muscle tone. Those with predominantly ascending projections discharge in association with fast, gamma electroencephalogram (EEG) activity and cease firing with slow, delta activity to be active during both wakefulness and rapid eye movement sleep (W/REM, filled red symbols); they include neurons which release glutamate (Glu, diamonds) or acetylcholine (ACh, circles) (see Figures 9.2 and 9.3). Those with more diffuse or descending projections discharge in association with behavioral arousal and electromyogram (EMG) activity and cease firing with muscle atonia to be active during W and silent during REM (W, empty red symbols); they include neurons which release glutamate (Glu, diamonds), noradrenaline (norepinephrine) (NA, square), serotonin (Ser, star), histamine (HA, cross) or orexin (Orx, asterisk) (see Figure 9.4). Neurons which are active during sleep (blue or aqua symbols) include cells with ascending projections toward the cortex, which dampen fast cortical activity, and those with descending projections toward the hypothalamus, brainstem, or spinal cord, which diminish behavioral arousal and muscle tone. Those with projections to the cortex or local area discharge in association with slow EEG activity during slow-wave sleep (SWS, blue triangle; see Figure 9.3); those with descending projections discharge in association with decreasing muscle tone and EMG (SWS/REM, aqua triangles; see Figure 9.3). They include particular GABAergic neurons in the basal forebrain and preoptic area that bear α2-adrenergic receptors and are thereby inhibited by NA. Also shown are GABAergic neurons in the pontomesencephalic tegmentum, which can inhibit local reticular or monoaminergic neurons, and GABAergic neurons (together with glycinergic neurons, not shown) in the ventral medullary reticular formation that project directly to the spinal cord where they can inhibit neck and other motor neurons during sleep. BF, basal forebrain; CB, cerebellum; Cx, cortex; DR, dorsal raphe; GABA, gamma-aminobutyric acid; LC, locus coeruleus nucleus; LDT, laterodorsal tegmental nucleus; Mes, mesencephalon; PH, posterior hypothalamus; POA, preoptic area; RF, reticular formation; SC, spinal cord; Th, thalamus; TM, tuberomammillary nucleus; VTA, ventral tegmental area. (Adapted from Jones (2005).)

Discharge of a cholinergic basal forebrain neuron across sleep–wake states. Record of a neuron labeled by juxtacellular technique with Neurobiotin (Nb) and identified by immunohistochemistry for choline acetyltransferase (ChAT) as cholinergic in the magnocellular preoptic nucleus (MCPO) of the rat. As evident in 10-second traces (above), the unit fired during aW, virtually ceased firing during SWS, resumed firing during tPS, and discharged maximally during PS. As evident in expanded 0.5-second traces (below), the unit discharged in rhythmic bursts of spikes with theta EEG activity that was present intermittently during periods of aW, toward the end of tPS, and continuously during PS. aW, active wake; EMG, electromyogram; EEG, electroencephalogram; PF, prefrontal cortex; RS, retrosplenial cortex; SWS, slow-wave sleep; tPS, transition to paradoxical sleep; PS, paradoxical sleep. Bar for horizontal scale: 1 second. Bar for vertical scales: 1 mV for EEG/EMG and 1.5 mV for unit. (Reprinted with permission from Lee et al. (2005b).)

Sleep–wake-related electroencephalogram (EEG)/electromyogram (EMG) and unit activity of basal forebrain neurons in the rat. Normalized average gamma power (A), delta power (B), and EMG amplitude (C) across all sleep–wake stages in the rat. (D–G) Normalized average unit spike rate for basal forebrain cell groups. In D, waking (W)/paradoxical sleep (PS)-active cells, whose discharge is positively correlated with gamma EEG activity and negatively correlated with delta EEG activity (including putative cholinergic cells represented in Figure 9.1 in the human brain as W/REM cells, circles). E shows W-active cells whose discharge is positively correlated with EMG amplitude and which fire maximally during W (including putative glutamatergic cells represented in Figure 9.1, diamonds). F shows SWS-active cells whose discharge is negatively correlated with gamma and positively correlated with delta EEG activity and which fire maximally during SWS (including putative GABAergic neurons represented in Figure 9.1, triangles). G shows SWS/PS-active cells whose discharge is negatively correlated with EMG amplitude and which fire at progressively higher rates during SWS through PS (including putative GABAergic neurons represented in Figure 9.1 as SWS/REM cells, triangles). aW, active wake; qW, quiet wake; tSWS, transition to slow-wave sleep; SWS, slow-wave sleep; tPS, transition to paradoxical sleep; PS, paradoxical sleep. (Reprinted with permission from Jones, (2005).)

Discharge of an Orx neuron across sleep–wake states. Record of a neuron labeled by juxtacellular technique with Neurobiotin (Nb) and identified by immunohistochemistry for Orx in the rat. As evident in 10-second traces (above), the unit fired during wakefulness (A) and was virtually silent during slow-wave sleep (B), transition to paradoxical sleep (C), and paradoxical sleep (D). As evident in an expanded trace (of approximately 4 seconds, below), the unit discharged during active wake (aW) and increased firing phasically in association with increases in muscle tone seen on the EMG. aW, active wake; EEG, electroencephalogram; EMG. electromyogram; PF, prefrontal cortex; PS, paradoxical sleep; RS, retrosplenial cortex; SWS, slow-wave sleep; tPS, transition to paradoxical sleep; W, wake. Horizontal scale bars: 1 second. Vertical scale bar: 1 mV for EEG, 0.5 mV for EMG, and 2 mV for unit. (Reprinted with permission from Lee et al. (2005a).
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