外侧缰核与外侧视前区在睡眠觉醒调节中的功能联系
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摘要
睡眠是生物机体的一种重要本能行为,当今社会睡眠障碍及由睡眠障碍引起的精神疾病的发生率逐年上升,造成严重的社会危害,因此睡眠研究已成为当今社会及医学界研究的热点。位于丘脑的缰核,尤其是外侧缰核在中枢与许多睡眠相关中枢有联系,其中与下丘脑外侧视前区(LPO)的纤维联系更为密切。近年关于外侧缰核的功能引起国外一些学者的关注,尤其是它在抑郁症发病中的关键作用引起人们极大的兴趣和重视,而抑郁症最典型的临床表现是睡眠障碍,两者互为因果。因此,外侧缰核可能参与睡眠觉醒周期的调节,并且可能是通过外侧视前区—外侧缰核通路发挥调节睡眠周期的作用。
本实验应用动物脑电记录方法,首先观察化学损毁外侧缰核后的大鼠在正常条件下、24小时持续光照(LL)或持续黑暗(DD)条件下24小时睡眠觉醒周期的变化,并比较6小时睡眠剥夺动物在外侧缰核缺失后24小时睡眠觉醒周期改变,结果显示:无论正常大鼠,还是6小时睡眠剥夺动物,损毁外侧缰核后,都表现觉醒时间增加,提示外侧缰核具有促进睡眠调节作用,不仅调节睡眠的总量,而且调节睡眠的深度和倾向性,并且外侧缰核也参与睡眠剥夺后的睡眠恢复过程。LL条件下,正常大鼠觉醒时间降低而非快眼动睡眠(NREMS)增加,但损毁外侧缰核的大鼠觉醒时间没有变化;而DD条件下,损毁组大鼠与正常组大鼠睡眠觉醒时间没有差异,提示外界光可诱导外侧缰核发挥调节睡眠的作用。
LPO具有促睡眠作用已被证实,我们进一步应用神经元细胞外记录、免疫组织化学测定等记录方法,研究外侧缰核对LPO促睡眠作用的影响及可能涉及的机制,将会从另一角度理解外侧缰核在睡眠觉醒调节中的作用。结果显示:外侧缰核神经元放电对LPO内电刺激、谷氨酸和γ-氨基丁酸分别化学兴奋和抑制、睡眠物质腺苷兴奋分别表现兴奋、兴奋和抑制、兴奋反应,而兴奋外侧缰核,LPO表现抑制效应;3小时睡眠剥夺组大鼠LPO内Fos+细胞数明显增加,并且可见来自于外侧缰核的逆行标记CTb+细胞与之重叠。这些结果表明,外侧缰核与LPO之间存在密切的功能联系,并且两者之间相互影响,构成一个负反馈环路参与睡眠的调节,这为中枢睡眠觉醒周期调节系统勾画一条新的途径。
Sleep is a very important life situation about one-third of our life time. In recent years, with the accelerated pace of life, sleep deprivation at night operation, and a variety of occupational groups are common, and sleep-related diseases such as insomnia, sleep disorders, the incidence of depression has increased year by year, so the current study has been more and more focus on the sleep mechanism in medical profession, biology and society.
Many central nucleus regulated sleep-wake cycle, such as lateral preoptic area, suprachiasmatic nucleus, raphe nuclei, pineal gland and other parts, and they involved in the sleep center to maintain homeostasis of sleep. Habenular nucleus sites in the epithalamus and have fibers linked with these sleep center. In recent years, the function of the lateral habenular nucleus caused a number of the attention of scholars, especially in the effects of LHb in the pathogenesis of depression. And the most typical clinical manifestation of depression is a sleep disorder. Therefore, the lateral habenular nucleus may be involved in the regulation of sleep-wakefulness cycle. Lateral habenular received projection fibers from the retinal ganglion directly, which showed its own circadian rhythm activities. Furthermore the lateral habenular nucleus contains neurotransmitter, neuromodulator and its receptor, which is involved in regulation of sleep homestatstic. Lateral habenular nucleus received efferent from lateral preoptic area fiber which is a sleep-promoting center. Therefore, we speculated that the lateral habenular play an important role in the impact of sleep through the lateral preoptic area - lateral habenular nucleus way.
To solve the above problem, first of all, we used EEG recording technique to study whether lateral habenular nucleus regulating sleep and affecting which parts of sleep as well as sleep deprivation.Then we used electrophysiological methods to explore the function between the lateral habenular nucleus and lateral preoptic area.Further immunohistochemical method was used to determine the lateral preoptic area - the lateral habenular nucleus pathway in the role of sleep-wake cycle.
The research results showed that:
(1) The effects of LHb regulating sleep-wake cycle:
Following lesion of the LHb and compared with the rats before lesion, Wake time was increased (P<0.001). Meanwhile, NREMS was decreased (P<0.001). REM was decreased from (P>0.05), but showed no significantly difference. In lesion rat, wakefulness was increased from 213.02±8.5min to269.31±23.07min (t=-3.20 , P<0.05)and NREMS was decreased from 458.56±9.04min to 396.86±22.03min (t=4.262,P<0.01)during the light-on period. Whereas REM showed no significantly difference. In lesion rat, wakefulness was increased from 451.28±24.92 min to 519.67±13.81min (t=-2.583, P<0.05) during the light-off period. NREMS and REMS tended to decrease but this trend was not statistically significant.
(2)The effects of LHb in the phonic reaction:
In at 24 hours light continuous illumination, the wakfulness was reduced (P <0.05) and NREMS was increased (P <0.05), REMS was lower in normal rats (P> 0.05)compared with in the LD condition normal rats. But the lesion lateral habenular nucleus of rats showed no difference between in LD condition and DD condition (P> 0.05). The wake time and NREMS both the normal rats and lesion lateral habenular nucleus rats showed no significantly change both in LD and DD.
(3) The effects of LHb in sleep deprivation:
After the 6 hours of sleep deprivation, the wakefulness in lateral habenular rats was significantly greater than the normal rat group And detla activity in lesion rats was increased compared with normal rats during sleep deprivation time.It indicate that lateral habenular join in the depth of sleep (P <0.05). The theta activity of lesion rats was significantly lower than normal group (P <0.05) indicated the lateral habenular nucleus increased sleep tendency.
(4) The effects of microinjection of AD antagonist into LHb in regulating sleep: After microinjection of adenosine receptor antagonist into lateral habenular nucleus, NREMS was significantly reduced from 148.28±3.51min to 137.31± 5.33min (t = 3.267, P<0.05). wakefulness was increased from 65.29±4.77min to 76.19±5.3min (t =- 2.608, P<0.05). REMS showed no change from 25.83±3.73min to 26.17±1.66min.
(5) The electophysiological function between LHb and LPO:
Application of single-pulse stimulation and tetanic methods to stimulate the lateral preoptic area and record the lateral habenular nucleus discharge showed that the discharge in lateral habenular nucleus was increased. Return stimulate lateral habenular nucleus and record the discharge of lateral preoptic area neurons show that the firing rate of LPO was reduced. After electic-lesion of stria medullary then uesd single pulse stimulation lateral preoptic area to record lateral habenular neurons showes no change.
(6) The effects of GABA and Glu on the function between LHb and LPO:
Electrochemical studies have shown that the effect of microinjection Glu into LPO on the firing rate of neurons in LHb was excited. The firing rate of LPO was decreased after microinjection of Glu in LHb. The effect of microinjection GABA into LPO on the firing rate of neurons in LHb was inhibition. The firing rate of LPO was increased after microinjection of GABA in LHb.
(7) The effects of microinjection of AD and AD antagonist into LPO in the firing rates of LHb:
The firing rate of LHb was increased after microinjection of AD into LPO (from 4.67±0.71Hz to 6.22±0.86Hz, P<0.05). The firing rate of LHb was increased after microinjection of 8-PT into LHb was significantly increased from 3.63±0.57Hz to 6.11±1.08Hz (P<0.05).
(8) CTb and c-fos double staining:
Immunohistochemical staining showed that Fos + CTb + cell count was increased in three hours of sleep deprivation 114.1±2.49, compared to in rats normal sleep group 16.25±1.01 (P <0.001)
From the above results we can draw the following conclusions:
(1) Application of animal EEG recording showed, wakefulness was increased and NREMS was decreased in lesion rat during the 24 hours compared with normal rats. It suggest that lateral habenular nucleus may play a critical role in the regulation of total amount of sleep. In LL, the wakfulness was reduced and NREMS was increased in normal rats,but the lesion lateral habenular nucleus of rats showed no difference between in LD condition and DD condition. Both the normal rats and lesion lateral habenular nucleus rats showed no significantly change both in LD and DD. These suggest that external light may be a key role in LHb regulating sleep.Microinjection of adenosine antagonists into LHb can increase the wake time, show that the lateral habenular approved as the role of endogenous adenosine targets involved in the regulation of sleep homeostasis.
2. We used electrical stimulation, Glu and GABA, respectively, which chemical excited and inhibition LHb, sleep-promoting substance adenosine excited LPO.The effects of LHb neurons were excite, excitement and inhibition, excitatory response. While we excited LHb, LPO neurons showed inhibitory effect, suggesting that the function LHb and LPO was close contact and mutual influence, which may form a negative feedback loop involved in the regulation of sleep.
3. With six hours of sleep deprivation in animals, lesion of the LHb rats showed increased wake time, NREMS was increased and detla activity was increased, as well as the lower theta wave. Lateral habenular nucleus microinjection of retrograde tracer (CTb), after 3 hours of sleep deprivation showed more c-fos + and CTb + double-stained cells in LPO in the SD group compared with normal sleep group. It suggested that LPO - LHb participate in the regulation of sleep-wakefulness cycle, to increase of sleep depth and propensity, which is related to these two parts involved in the regulation of recovery sleep after sleep deprivation.
Our study found that the lateral habenular nucleus is a new center in sleep-promoting, not only regulating the total amount of sleep and regulating depth and propensity. More importantly, defined the LPO– LHb constitute a negative feedback pathway in regulating sleep, but it is not very clear that mechanisms between the two nuclei regulation sleep. In the future Research we will carry out further exploration.
本实验应用动物脑电记录方法,首先观察化学损毁外侧缰核后的大鼠在正常条件下、24小时持续光照(LL)或持续黑暗(DD)条件下24小时睡眠觉醒周期的变化,并比较6小时睡眠剥夺动物在外侧缰核缺失后24小时睡眠觉醒周期改变,结果显示:无论正常大鼠,还是6小时睡眠剥夺动物,损毁外侧缰核后,都表现觉醒时间增加,提示外侧缰核具有促进睡眠调节作用,不仅调节睡眠的总量,而且调节睡眠的深度和倾向性,并且外侧缰核也参与睡眠剥夺后的睡眠恢复过程。LL条件下,正常大鼠觉醒时间降低而非快眼动睡眠(NREMS)增加,但损毁外侧缰核的大鼠觉醒时间没有变化;而DD条件下,损毁组大鼠与正常组大鼠睡眠觉醒时间没有差异,提示外界光可诱导外侧缰核发挥调节睡眠的作用。
LPO具有促睡眠作用已被证实,我们进一步应用神经元细胞外记录、免疫组织化学测定等记录方法,研究外侧缰核对LPO促睡眠作用的影响及可能涉及的机制,将会从另一角度理解外侧缰核在睡眠觉醒调节中的作用。结果显示:外侧缰核神经元放电对LPO内电刺激、谷氨酸和γ-氨基丁酸分别化学兴奋和抑制、睡眠物质腺苷兴奋分别表现兴奋、兴奋和抑制、兴奋反应,而兴奋外侧缰核,LPO表现抑制效应;3小时睡眠剥夺组大鼠LPO内Fos+细胞数明显增加,并且可见来自于外侧缰核的逆行标记CTb+细胞与之重叠。这些结果表明,外侧缰核与LPO之间存在密切的功能联系,并且两者之间相互影响,构成一个负反馈环路参与睡眠的调节,这为中枢睡眠觉醒周期调节系统勾画一条新的途径。
Sleep is a very important life situation about one-third of our life time. In recent years, with the accelerated pace of life, sleep deprivation at night operation, and a variety of occupational groups are common, and sleep-related diseases such as insomnia, sleep disorders, the incidence of depression has increased year by year, so the current study has been more and more focus on the sleep mechanism in medical profession, biology and society.
Many central nucleus regulated sleep-wake cycle, such as lateral preoptic area, suprachiasmatic nucleus, raphe nuclei, pineal gland and other parts, and they involved in the sleep center to maintain homeostasis of sleep. Habenular nucleus sites in the epithalamus and have fibers linked with these sleep center. In recent years, the function of the lateral habenular nucleus caused a number of the attention of scholars, especially in the effects of LHb in the pathogenesis of depression. And the most typical clinical manifestation of depression is a sleep disorder. Therefore, the lateral habenular nucleus may be involved in the regulation of sleep-wakefulness cycle. Lateral habenular received projection fibers from the retinal ganglion directly, which showed its own circadian rhythm activities. Furthermore the lateral habenular nucleus contains neurotransmitter, neuromodulator and its receptor, which is involved in regulation of sleep homestatstic. Lateral habenular nucleus received efferent from lateral preoptic area fiber which is a sleep-promoting center. Therefore, we speculated that the lateral habenular play an important role in the impact of sleep through the lateral preoptic area - lateral habenular nucleus way.
To solve the above problem, first of all, we used EEG recording technique to study whether lateral habenular nucleus regulating sleep and affecting which parts of sleep as well as sleep deprivation.Then we used electrophysiological methods to explore the function between the lateral habenular nucleus and lateral preoptic area.Further immunohistochemical method was used to determine the lateral preoptic area - the lateral habenular nucleus pathway in the role of sleep-wake cycle.
The research results showed that:
(1) The effects of LHb regulating sleep-wake cycle:
Following lesion of the LHb and compared with the rats before lesion, Wake time was increased (P<0.001). Meanwhile, NREMS was decreased (P<0.001). REM was decreased from (P>0.05), but showed no significantly difference. In lesion rat, wakefulness was increased from 213.02±8.5min to269.31±23.07min (t=-3.20 , P<0.05)and NREMS was decreased from 458.56±9.04min to 396.86±22.03min (t=4.262,P<0.01)during the light-on period. Whereas REM showed no significantly difference. In lesion rat, wakefulness was increased from 451.28±24.92 min to 519.67±13.81min (t=-2.583, P<0.05) during the light-off period. NREMS and REMS tended to decrease but this trend was not statistically significant.
(2)The effects of LHb in the phonic reaction:
In at 24 hours light continuous illumination, the wakfulness was reduced (P <0.05) and NREMS was increased (P <0.05), REMS was lower in normal rats (P> 0.05)compared with in the LD condition normal rats. But the lesion lateral habenular nucleus of rats showed no difference between in LD condition and DD condition (P> 0.05). The wake time and NREMS both the normal rats and lesion lateral habenular nucleus rats showed no significantly change both in LD and DD.
(3) The effects of LHb in sleep deprivation:
After the 6 hours of sleep deprivation, the wakefulness in lateral habenular rats was significantly greater than the normal rat group And detla activity in lesion rats was increased compared with normal rats during sleep deprivation time.It indicate that lateral habenular join in the depth of sleep (P <0.05). The theta activity of lesion rats was significantly lower than normal group (P <0.05) indicated the lateral habenular nucleus increased sleep tendency.
(4) The effects of microinjection of AD antagonist into LHb in regulating sleep: After microinjection of adenosine receptor antagonist into lateral habenular nucleus, NREMS was significantly reduced from 148.28±3.51min to 137.31± 5.33min (t = 3.267, P<0.05). wakefulness was increased from 65.29±4.77min to 76.19±5.3min (t =- 2.608, P<0.05). REMS showed no change from 25.83±3.73min to 26.17±1.66min.
(5) The electophysiological function between LHb and LPO:
Application of single-pulse stimulation and tetanic methods to stimulate the lateral preoptic area and record the lateral habenular nucleus discharge showed that the discharge in lateral habenular nucleus was increased. Return stimulate lateral habenular nucleus and record the discharge of lateral preoptic area neurons show that the firing rate of LPO was reduced. After electic-lesion of stria medullary then uesd single pulse stimulation lateral preoptic area to record lateral habenular neurons showes no change.
(6) The effects of GABA and Glu on the function between LHb and LPO:
Electrochemical studies have shown that the effect of microinjection Glu into LPO on the firing rate of neurons in LHb was excited. The firing rate of LPO was decreased after microinjection of Glu in LHb. The effect of microinjection GABA into LPO on the firing rate of neurons in LHb was inhibition. The firing rate of LPO was increased after microinjection of GABA in LHb.
(7) The effects of microinjection of AD and AD antagonist into LPO in the firing rates of LHb:
The firing rate of LHb was increased after microinjection of AD into LPO (from 4.67±0.71Hz to 6.22±0.86Hz, P<0.05). The firing rate of LHb was increased after microinjection of 8-PT into LHb was significantly increased from 3.63±0.57Hz to 6.11±1.08Hz (P<0.05).
(8) CTb and c-fos double staining:
Immunohistochemical staining showed that Fos + CTb + cell count was increased in three hours of sleep deprivation 114.1±2.49, compared to in rats normal sleep group 16.25±1.01 (P <0.001)
From the above results we can draw the following conclusions:
(1) Application of animal EEG recording showed, wakefulness was increased and NREMS was decreased in lesion rat during the 24 hours compared with normal rats. It suggest that lateral habenular nucleus may play a critical role in the regulation of total amount of sleep. In LL, the wakfulness was reduced and NREMS was increased in normal rats,but the lesion lateral habenular nucleus of rats showed no difference between in LD condition and DD condition. Both the normal rats and lesion lateral habenular nucleus rats showed no significantly change both in LD and DD. These suggest that external light may be a key role in LHb regulating sleep.Microinjection of adenosine antagonists into LHb can increase the wake time, show that the lateral habenular approved as the role of endogenous adenosine targets involved in the regulation of sleep homeostasis.
2. We used electrical stimulation, Glu and GABA, respectively, which chemical excited and inhibition LHb, sleep-promoting substance adenosine excited LPO.The effects of LHb neurons were excite, excitement and inhibition, excitatory response. While we excited LHb, LPO neurons showed inhibitory effect, suggesting that the function LHb and LPO was close contact and mutual influence, which may form a negative feedback loop involved in the regulation of sleep.
3. With six hours of sleep deprivation in animals, lesion of the LHb rats showed increased wake time, NREMS was increased and detla activity was increased, as well as the lower theta wave. Lateral habenular nucleus microinjection of retrograde tracer (CTb), after 3 hours of sleep deprivation showed more c-fos + and CTb + double-stained cells in LPO in the SD group compared with normal sleep group. It suggested that LPO - LHb participate in the regulation of sleep-wakefulness cycle, to increase of sleep depth and propensity, which is related to these two parts involved in the regulation of recovery sleep after sleep deprivation.
Our study found that the lateral habenular nucleus is a new center in sleep-promoting, not only regulating the total amount of sleep and regulating depth and propensity. More importantly, defined the LPO– LHb constitute a negative feedback pathway in regulating sleep, but it is not very clear that mechanisms between the two nuclei regulation sleep. In the future Research we will carry out further exploration.
引文
[1] SATEIA MJ. Update on sleep and psychiatric disorders [J]. Chest, 2009, 135 (5): 1370-1379.
[2] PICKERING TG, DEVEREUX RB, JAMES GD, et al. Environmental infuences on blood pressure and the role of job train [J]. J Hypertens Suppl, 1996, 14 (5): 179-185.
[3] Sleep Disorders and Sleep Deprivation: an Unmet Public Health Problem. Institute of Medicine of the National Academies[C]. The National Academies Press, Washington DC, 2006.
[4] VON ECONOMO C. Sleep as a problem of localization [J]. J Nerv Ment Dis, 1930, 71: 249–259.
[5]姚泰.生理学(第五版)[M].北京:人民卫生出版社,2000:333.
[6] MONTI JM, JANTOS H. The roles of dopamine and serotonin, and of their receptors, in regulating sleep and waking [J]. Prog Brain Res, 2008, 172: 625-646.
[7] SZYMUSIAK R, McGINTY D. Hypothalamic regulation of sleep and arousal [J]. Ann N Y Acad Sci, 2008, 1129: 275-286.
[8] CESPUGLIO R, SARDA N, GHARIB A, et al. Voltammetric detection of the release of 5-hydroxyindole Compounds throughout the sleep-waking cycle of the rat [J]. Exp Brain Res, 1990, 80 (1): 121-128.
[9] PORTAS CM, BJORVATN B, FAGERLAND S. Detection of extracellular levels of serotonin in dorsal raphe nucleus and frontal cortex over the sleep/wake cycle in the freely moving rat [J]. Neuronscience, 1998, 83 (3): 807-814.
[10] JOUVET M. Sleep and neurotransmitters [J]. Rev Med Interne, 1996, 17: 18-23.
[11] STERIADE M, MCCARLEY RW. Brain control of sleep and wakefulness [M]. New York: Kluwer Academic Press, 2005.
[12] BERRIDGE CW, WATERHOUSE BD. The locus coeruleus-noradrenergic system: modulation of behavioral state and state-dependent cognitive processes. [J]. Brain Res Brain Res Rev, 2003, 42 (1): 33-84.
[13] THANKCHAN S, KAUR S, SHIROMANI PJ. Activity of pontine neurons during sleep and cataplexy in hypocretin knock-out mice. [J]. J Neurosci, 2009, 29 (5): 1580-1585.
[14] KOYAMA Y, TAKAHASHI K, KODAMA T, et al. State-dependent activity of neurons in the perifornical hypothalamic area during sleep and waking [J]. Neuroscience, 2003, 119 (4): 1209-1219.
[15] FRONCZEK R, BAUMANN CR, LAMMERS GJ, et al. Hypocretin/orexin disturbances in neurological disorders [J]. Sleep Medicine Reviews, 2009, 13 (1): 9-22.
[16] NAUTA WJH. Hypothalamic regulation of sleep in rats [J]. An experimentalstudy. J Neurophysiol, 1946, 9: 285–316.
[17] McGINTY D, STERMAN MB. Sleep suppression after basal forebrain lesions in the cat [J]. Science, 1968, 160: 1253–1255.
[18] SZYMUSIAK R, SATINOFF E. Ambient temperature-dependence of sleep disturbances produced by basal forebrain damage in rats [J]. Brain Res Bull, 1984, 12(3): 295–305.
[19] GONG H, SZYMUSIAK R, KING J, et al. Sleep-related c-Fos protein expression in the preoptic hypothalamus: effects of ambient warming [J]. Am J Physiol Regul Integr Comp Physiol, 2000, 279 (6): 2079-2088.
[20] SZYMUSIAK R, ALAM N, STEININGER TL, et al. Sleep-waking discharge patterns of ventrolateral preoptic/anterior hypothalamic neurons in rats [J]. Brain Res, 1998, 803 (1-2): 178–188.
[21] GONG H, McGINTY D, GUZMAN-MARIN R, et al. Activation of c-Fos in GABAergic neurones in the preoptic area during sleep and in response to sleep deprivation [J]. J Physiol, 2004, 556(Pt3): 935–946.
[22] PARAMENTIER R, KOLBAEV S, KLYUCH BP, et al. Excitation of histaminergic tuberomamillary neurons by thyrotropin-releasing hormone [J]. J Neurosci. 2009, 8, 29(14):4471-4483.
[23] MITANI A, ITO K, HALLANGER AE, et al. Cholinergic projections from the laterodorsal and pedunculopontine tegmental nuclei to the pontine gigantocellular tegmental field in the cat [J]. Brain Res, 1988, 451(1-2): 397–402.
[24] GNADT JW, PEGRAM GV. Cholinergic brainstem mechanisms of REM sleep in the rat [J]. Brain Res, 1986, 384 (1):29–41.
[25] STERIADE M, McCARLEY RW. Brain control of sleep and wakefulness.New York: Kluwer Academic Press, 2005.
[26] McCARLEY RW. Mechanisms and models of REM sleep control [J]. Archives Ital de Biol, 2004, 142: 429–468.
[27] FULLER PM, SAPER CB, LU J. The pontine REM switch: past and present [J]. J Physiol, 2007, 584(Pt 3): 735-741.
[28] GREENE RW, GERBER U, McCARLEY RW. Cholinergic activation of medial pontine reticular formation neurons in vitro [J]. Brain Res, 1989, 476: 154–159.
[29] FORNAL C, AUERBACH S, JACOBS BL. Activity of serotonin- containing neurons in nucleus raphe magnus in freely moving cats [J]. Exp Neurol, 1985,88(3): 590–608.
[30] LEVINE ES, JACOBS BL. Neurochemical afferents controlling theactivity of serotonergic neurons in the dorsal raphe nucleus: microiontophoretic studies in the awake cat [J]. J Neurosci, 1992, 12:4037–4044.
[31] GERVASONI D, DARRACQ L, FORT P, et al. Electrophysiological evidence that noradrenergic neurons of the rat locus coeruleus are tonically inhibited by GABA during sleep [J]. Eur J Neurosci, 1998, 10:964–970.
[32] JONES IS. The organization of central cholinergic systems and their functional importance in sleep–waking states[J]. Prog Brain Res, 1993, 98: 61–71.
[33] TORTEROLO P, MORALES FR, CHASE MH. GABAergic mechanisms in the pedunculopontine tegmental nucleus of the cat promote active (REM) sleep [J]. Brain Res, 2002, 944(1-2): 1-9.
[34] RADULOVACKI M. Role of adenosine in sleep in rats [J]. Rev Clin Basic Pharmacol, 1985, 5(3-4):327–339.
[35] TICHO SR, RADULOVACKI M. Role of AD in sleep and temperature regulation in the preoptic area of rats [J]. Pharmacol Biochem Behav, 1991, 40(1):33–40.
[36] PHILLIS JW, WU PH. The role of adenosine and its nucleotide in central synaptic transmission [J]. Prog Neurobiol, 1981, 16(3-4): 187–239.
[37] PHILLIS J.W., WALTER G.A., O’REGAN M.H., et al. Adenosine in the control of cerebral circulation [J]. Cerebrovasc. Brain Metab. Rev, 1989, 1: 26–54.
[38] TICHO SR, RADULOVACKI M. Role of adenosine in sleep and temperature regulation in the preoptic area of rats [J]. Pharmacol Biochem Behav, 1991, 40(1):33-40.
[39] FREDHOLM BB. Astra Award Lecture. Adenosine, adenosine receptors and the actions of caffeine [J]. Pharmacol Toxicol, 1995, 76(2):93-101.
[40] FREDHOLM BB, ARSLAN G, HALLDNER L, et al. Structure and function of adenosine receptors [J]. Naunyn Schmiedebergs Arch Pharmacol, 2000, 362(4-5): 364–374.
[41] KLOTZ KN. Adenosine receptors and their ligands [J]. Naunyn Schmiedebergs Arch Pharmacol, 2000, 362(4-5):382–391.
[42] PORKKA-HEISKANEN T, STRECKER RE, THAKKAR M, et al. Adenosine: a mediator of the sleep-inducing effects of prolonged wakefulness [J]. Science,1997, 276(5316): 1265– 1268.
[43] PORKKA-HEISKANEN T, STRECKER RE, McCARLEY RW. Brain site specificity of extracellular adenosine concentration changes during sleep deprivation and spontaneous sleep: an in vivo microdialysis study [J]. Neuroscience 2000, in press.
[44] RAINNIE DG, GRUNZE HC, McCARLEY RW, et al. Adenosine inhibition of mesopontine cholinergic neurons: implications for EEG arousal [J]. Science, 1994, 263:689–692.
[45] ROBERT E. STRECKER, STEPHEN MORAIRTY, MAHESH M THAKKER, et al .Adenosinergic modulation of basal forebrain and preoptic/anterior hypothalamic neuronal activity in the control of behavioral state[J]. Behavioural Brain Research, 2000, 115(2):183-204.
[46] MATSUMURA H, NAKAJIMA T, OSAKA T, et al. Prostaglandin D2-sensitive, sleep promoting zone defined in the ventral surface of the rostral basal forebrain [J]. Proc Natl Acad Sci USA, 1994, 91(25):11998–12002.
[47] SATOH S, MATSUMURA H, SUZUKI F, et al. Promotion of sleep mediated by the A2a-adenosine receptor and possible involvement of this receptor in the sleep induced by prostaglandin D2 in rats [J]. Proc Natl Acad Sci U S A., 1996, 93(12):5980-5984.
[48] MIZOGUCHI A, EGUCHI N, KIMURA K, et al. Dominant localization of prostaglandin D receptors on arachnoid trabecular cells in mouse basal forebrain and their involvement in the regulation of non-rapid eye movement sleep [J]. Proc Natl Acad Sci USA, 2001, 98(25):11674–11679.
[49] MORAIRTY S, RAINNIE D, McCARLEY R, et al. Disinhibition of ventrolateral preoptic area sleep-active neurons by adenosine: a new mechanism for sleep promotion [J]. Neuroscience, 2004, 123(2):451-457.
[50] ERIC E. ABRAHAMSON, ROBERT Y. MOORE. Suprachiasmatic nucleus in the mouse: retinal innervation, intrinsic organization and efferent projections [J]. Brain Research, 2001, 916(1-2): 172–191.
[51] LESAUTER J, LEHMAN MN, SILVER R. Restoration of circadian rhythmicity by transplants of SCN "micropunches"[J]. J Biol Rhythms, 1996, 11(2):163-171.
[52] IBUKA N, KAWAMURA H. Loss of circadian rhythm in sleepwakefulness cycle in the rat by suprachiasmatic nucleus lesions [J]. Brain Res, 1975, 96(1):76-81.
[53] MOORE RY, EICHLER VB. Loss of a circadian adrenal corticosterone rhythm following suprachiasmatic lesions in the rat [J]. Brain Res, 1972, 42(1):201-206.
[54]周先举,俞光第,印其章.哺乳动物昼夜节律组构中的下丘脑视交叉上核和松果腺.生理科学进展[J],2001,32(2):116-120.
[55] CHALLET E, CALDELAS I, GRAFF C, et al. Synchronization of the molecular clockwork by light- and food-related cues in mammals [J]. Biol Chem, 2003, 384(5):711-719.
[56] ZHAO H., RUSAK B. Circadian fring-rate rhythms and light responses of rat habenular nucleus neurons in vivo and in vitro [J]. Neuroscience, 2005, 132: 519–528.
[57] MATTHEWS-FELTON T, CORODIMAS KP, ROSENBLATT JS, et al. Lateral habenula neurons are necessary for the hormonal onset of maternal behavior and for the display of postpartum estrus in naturally parturient female rats [J]. Behav Neurosci, 1995, 109(6): 1172–1188.
[58] GOLDSTEIN RA. GABAergic habenulo-raphe pathway mediation of the hypnogenic effects of vasotocin in the cat [J]. Neuroscience, 1983, 10(3):941–945.
[59] GALLAGHER M., CHIBA AA. The amygdala and emotion [J]. Neurobiology, 1996, 6(2):221-227.
[60]王绍.缰核与奖赏和稳态[J].生理科学进展,2008,4(39): 292-296.
[61] HERKENHAM M., NAUTA W.J.H. Efferent connections of the habenular nuclei in the rat [J]. Journal of Comparative Neurology, 1979, 187(1): 19–47.
[62] ANDRES KH, VON DURING M, VEH RW. Subnuclear organization of the rat habenular complexes [J].The Journal of Comparative Neurology, 1999, 407(1): 130–150.
[63] KIM U, CHANG SY. Dendritic morphology, local circuitry and intrinsic electrophysiology of neurons in the rat medial and lateral habenular nuclei of the epithalamus [J]. J COMP NEURO, 2005, 483 (2): 236-250.
[64] ARAKI M, MCGEER PL, MCGEER, EG. Retrograde HRP tracing combined with a pharmacohistochemical method for GABA transaminase for the identification of presumptive GABAergic projections to the habenula [J]. Brain Research, 1984, 304(2): 271–277.
[65] GARLAND JC, MOGENSON GJ. An electrophysiological study of convergenceof entopeduncular and lateral preoptic inputs on lateral habenular neurons projecting to the midbrain [J]. Brain Research, 1983, 263: 33–41.
[66] VINCENT SR, KIMURA H, MCGEER EG. A histochemical study of GABA-transaminase in the efferents of the pallidum [J]. Brain Research, 1982, 241(1): 162–165.
[67] NAGY JI, CARTER DA, LEHMANN J, et al. Evidence for a GABA-containing projection from the entopedoncular nucleus to the lateral habenula in the rat [J]. Brain Research, 1978, 145(2):360–364.
[68] CHANG S, KIM U. Ionic mechanism of long-lasting discharges of action potentials triggered by membrane hyperpolarization in the medial lateral habenula [J]. Journal of Neuroscience, 2004, 24: 2172–2181.
[69] MORIIZUMI T, HATTORI T. Choline acetyltransferase- immunoreactive neurons in the rat entopedoncular nucleus [J]. Neuroscience, 1992, 46(3):721–728.
[70] VERTES RP, FORTIN WJ, CRANE AM. Projections of the median raphe′nucleus in the rat [J]. Journal of Comparative Neurology, 1999, 407: 555–582.
[71] QU T, DONG K, SUGIOKA K, et al. Demonstration of direct input from the retina to the lateral habenular nucleus in the albino rat [J]. Brain Research, 1996, 709(2): 251–258.
[72] KAWAJA MD, FLUMERFEIL BA., HRYCYSHYN AM. Synaptic organization of septal projections in the rat medial habenula: a wheat germ agglutinin-horseradish peroxidase and immunohistochemical study [J]. Synapse, 1990, 6(1): 45–54.
[73] HERKENHAM M, NAUTA WJH. Afferent connections of the habenular nuclei in the rat. A horseradish peroxidase study, with a note on the fiber-of-passage problem [J]. Journal of Comparative Neurology, 1977, 173(1): 123–146.
[74] LECOURTIER L, KELLY PH. A conductor hidden in the orchestra? Role of the habenular complex inmonoamine transmission and cognition [J]. Neurosci Biobehav Rev, 2007, 31(5):658-672.
[75] HIKOSAKA O, SESACK SR, LECOURTIER L, et al. Habenula: crossroad between the basal ganglia and the limbic system [J]. J Neurosci, 2008, 28(46):11825-11829.
[76] GOLDSTEIN R. Vasotocin-like activity of cerebrospinal fluid induced byhabenular lesions in cats [J]. Neuropeptides, 1985, 6(4):303-310.
[77] MATTHEWS-FELTON T, LINTON L, ROSENBLATT JS, et al. First and second order maternal behavior related afferents of the lateral habenula [J]. Neuroreport, 1999, 10(4):883–887.
[78] SZYMUSIAK R, STEININGER T, ALAM N, et al. Preoptic area sleep-regulating mechanisms [J]. Arch Ital Biol, 2001, 139(1-2):77-92.
[79] FERRARO G, MONTALBANO ME, SARDO P, et al. Lateral habenula and hippocampus: a complex interaction raphe cells mediated [J]. Journal of Neural Transmission, 1997, 104(6-7): 615–631.
[80] ULLSPERGERM, YVES VON CRAMON D. The error monitoring using external feedback: specific roles of the habenular complex, the reward system, and the cingulate mot or area revealed by functi onal magnetic resonance imaging [J]. J Neurosci, 2003, 23: 4308-4314.
[81] JI H, SHEPARD PD. Lateral Habenula Stimulati on Inhibits Rat Midbrain Dopamine Neurons through a GABAA Receptor Mediated Mechanism [J]. J Neurosci, 2007, 27: 6923～6930 .
[82] MATSUMOTO M, HIKOSAKA O. Lateral habenula as a source of negative reward signals in dopamine neurons [J]. Nature, 2007, 447(7148): 1111-1115.
[83]赵华,王绍.分别兴奋大鼠内侧和外侧缰核引起痛阈的不同改变[J].生理学报,1995, 3(47): 292-296.
[84] YANG LM, HU B, XIA YH, et al. Lateral habenula lesions improve the behavioral response in depressed rats via increasing the serotonin level in dorsal raphe nucleus [J]. Behav Brain Res, 2008, 188(1):84-90.
[85] HAUN F, ECKENRODE TC, MURRAY M. Habenula and thalamus cell transplants restore normal sleep behaviors disrupted by denervation of the interpeduncular nucleus [J]. J Neurosci, 1992, 12(8):32.
[86] KELEIN DC, WELLER JL. Rapid light-induced decrease in pineal seroton N-acetyltransferase activity [J].Science, 1972, 177(48):532-533.
[87] LIU C, WEAVER DR, JIN X, et al. Molecular dissection of two distinct actions of melatonin on thesuprachiasmatic circadian clock [J].Neuron, 1997, 19:91-102.
[88] KOKKOLA T, LAITINEN JT. Melatonin receptor genes [J]. Ann Med, 1998, 30(1): 88-94.
[89]吴建群,王绍.缰核与GABA、乙酰胆碱[J].针刺研究,1989,1-2: 72- 73.
[90] CLAUDE GOTTESMANN. GABA mechanisms and sleep [J]. Neuroscience, 2002, 111(2): 231- 239.
[1] YANG LM, HU B, XIA YH, et al. Lateral habenula lesions improve the behavioral response in depressed rats via increasing the serotonin level in dorsal raphe nucleus [J]. Behav Brain Res, 2008, 188(1): 84-90.
[2] MORISSETTE MC, BOYE SM. Electrolytic lesions of the habenula attenuate brain stimulation reward [J]. Behav Brain Res, 2008, 187(1): 17-26.
[3] SARTORIUS A, HENN FA. Deep brain stimulation of the lateral habenula intreatment resistant major depression [J]. Med Hypotheses, 2007, 69(6): 1305-1308.
[4] ZHAO H, RUSAK B. Circadian firing-rate rhythms and light responses of rat habenular nucleus neurons in vivo and in vitro [J]. Neuroscience, 2005, 132(2): 519-528.
[5] KIM U, CHANG SY. Dendritic morphology, local circuitry and intrinsic electrophysiology of neurons in the rat medial and lateral habenular nuclei of the epithalamus [J]. J Comp Neurol, 2005, 483(2): 236-250.
[6] GOLDSTEIN R.A. GABAergic habenulo-raphe pathway mediation of the hypnogenic effects of vasotocin in cat [J]. Neuroscience, 1983, 10(3): 941-945.
[7] FLUMERFELT BA, CONTESTABILE A. Acetylcholinesterase Histochemistry of the habenulo-interpeduncular pathway in the rat and the effects of electrolytic and kainic acid lesions [J]. Anat Embryol (Berl), 1982, 163(4):435-446.
[8] IWAKIRI H, MATSUYAMA K, MORI S. Extracellular levels of serotonin in the medial pontine reticular formation in relation to sleep-wake cycle in cats: a microdialysis study [J]. Neurosci Res, 1993, 18(2):157-170.
[9] KOWSKI AB, GEISLER S, KRAUSS M, et al. Differential projections from subfields in the lateral preoptic area to the lateral habenular complex of the rat[J].J Comp Neurol, 2008, 507(4):1465-1478.
[10] ANDRES KH, VON DüRING M, VEH RW. Subnuclear organization of the rat habenular complexes [J]. J Comp Neurol, 1999, 407(1):130-150.
[11] GEISLER S, ANDRES KH, VEH RW. Morphologic and cytochemical criteria for the identification and delineation of individual subnuclei within the lateral habenular complex of the rat [J]. J Comp Neurol, 2003, 458(1):78-97.
[12] STAINES WA, YAMAMOTO T, DEWAR KM, et al. Distribution, morphology and habenular projections of adenosine deaminase-containing neurons in the septal area of rat [J]. Brain Res, 1988, 455(1):72-87.
[13] MATTEAU I, BOIRE D, PTITO M. Retinal projections in the cat: a cholera toxin B subunit study [J].Vis Neurosci, 2003, 20(5):481-493.
[14] SHIN TOKUNAGA, YASUHIRO TAKEDA, KAZUAKI SHINAMIYA, et al. Effects of some H1-antagonists on the sleep-Wake cycle in sleep-disturbed rats [J]. J Pharmacol Sci, 2007, 103(2):201-206.
[15] SHINOMIYA K, INOUE T, UTSU Y, et al. Effects of kava-kava extract on thesleep-wake cycle in sleep-disturbed rats [J]. Psychopharmacology (Berl), 2005, 180(3): 564-569.
[16] NECKELMANN D, URSIN R. Sleep stages and EEG power spectrum in relation to acoustical stimulus arousal threshold in the rat [J]. Sleep, 1993, 16(5): 467-477.
[17] FAULHABER J, STAIGER A, LANCEL M. The GABAA agonist THIP produces slow wave sleep and reduces spindling activity in NREM sleep in humans [J]. Psychopharmacology (Berl), 1997, 130(3):285–291.
[18] LANCEL M. The GABA (A) agonist THIP increases non-REM sleep and enhances non-REM sleep-specific delta activity in the rat during the dark period. Sleep, 1997, 20(12):1099–1104.
[19] ELLISON G. Stimulant-induced psychosis, the dopamine theory of schizophrenia, zand the habenula [J]. Brain Res Brian Res Rev, 1994, 19(2):223-239.
[20] HUSTON JP, HAAS HL, BOIX F, et al. Extracellular adenosine levels in neostriatum and hippocampusduring rest and activity periods of rats [J]. Neuroscience, 1996, 73(1):99-107.
[21] PORKKA-HEISKANEN T, STRECKER RE, MCCARLEY RW. Brain site-specificity of extracellular adenosine concentration changes during sleep deprivation and spontaneous sleep: an in vivo microdialysis study [J]. Neuroscience, 2000, 99(3):507-517.
[22] STRECKER RE, MORAIRTY S, THAKKAR MM, et al. Adenosinergic modulation of basal forebrain and preoptic/anterior hypothalamic neuronal activity in the control of behavioral state[J]. Behav Brain Res, 2000, 115(2):183-204.
[23] MURILLO-RODRIGUEZ E, BLANCO-CENTURION C, GERASHCHENKO D, et al. The diurnal rhythm of adenosine levels in the basal forebrain of young and old rats [J]. Neuroscience, 2004, 123(2):361-370.
[24] PORTAS CM, THAKKAR M, RAINNIE DG, et al. Role of adenosine in behavioral state modulation: a microdialysis study in the freely moving cat [J]. Neuroscience, 1997, 79(1):225-235.
[25] LUCAS EA, STERMAN MB. Effects of a forebrain lesion on the poly-cyclic sleep–wake cycle and sleep–wake patterns in the cat [J]. Exp Neurol, 1975, 46: 368–388.
[26] SZYMUSIAK R, McGINTY D. Sleep suppression following kainic acid-induced lesions of the basal forebrain [J]. Exp Neurol, 1986, 94:598–614.
[27] SALLANON M, DENOYER M, KITAHAMA K, et al. Long lasting insomnia induced by preoptic neuron lesions and its tran-sient reversal by muscimol injection into the posterior hypothalamus in the cat [J]. Neuroscience, 1989, 32: 669–683.
[28] SRIVIDYA R, MALLICK HN, KUMAR VM. Differences in the effects of medial and lateral preoptic lesions on thermoregulation and sleep in rats [J]. Neuroscience, 2006, 139(3):853-864.
[29] LECOURTIER L, NEIJT HC, KELLY PH. Habenula lesions cause impaired cognitive performance in rats: implications for schizophrenia [J]. Eur J Neurosci, 2004, 19(9):2551–2560.
[30] SHEPARD PD, HOLCOMB HH, GOLD JM. The presence of absence: Habenular regulation of dopamine neurons and the encoding of negative outcomes [J]. Schizophrenia Bull, 2006, 32(3):417–421.
[31] SHUMAKE J, EDWARDS E, GONZALEZ-LIMA F. Opposite metabolic changes in the habenula and ventral tegmental area of a genetic model of helpless behavior [J]. Brain Res, 2003, 963:274–281.
[32] FELTON T, LINTON L, ROSENBLATT JS, et al. First and second order maternal behavior related afferents of the lateral habenula [J]. Neuroreport, 1999, 10(4):883-887.
[33] BRISKI K, GILLEN E. Differential distribution of Fos expression within the male rat preoptic area and hypothalamus in response to physical vs. psychological stress [J]. Brain Res Bull, 2001, 55(3): 401-408.
[34] GOLDSTEIN R. Vasotocin-like activity of cerebrospinal fluid induced by habenular lesions in cats [J]. Neuropeptides, 1985, 6(4):303-310.
[35] GARLAND JC, MOGENSON GJ. An Electrophysiological Study of Convergence of Entopeduncular and LateralPreoptic Inputs on Lateral Habenular Neurons Projecting to the Midbrain [J]. Brain Research, 1983, 263(1):33-41.
[36] METHIPPARA MM, KUMAR S, ALAM MN, et al. Effects on sleep of microdialysis of adenosine A1 and A2A receptor analogs into the lateral preoptic area of rats [J]. Am J Physiol Regul Integr Comp Physiol, 2005, 89(6): 1715–1723.
[37] GVILIA I, FENG XU, MCGINTY D et al. Homeostatic Regulation of Sleep: A Role for Preoptic Area Neurons [J]. The Journal of Neuroscience, 2006, 26(37): 9426–9433.
[38] DéTáRI L, JUHáSZ G, KUKORELLI T. Firing properties of cat basal forebrain neurones during sleep-wakefulness cycle [J]. Electroencephalogr Clin Neurophysiol, 1984, 58(4):362-368.
[39] GVILIA I, TURNER A, MCGINTY D, et al. Preoptic area neurons and the homeostatic regulation of rapid eye movement sleep [J]. J Neurosci, 2006, 26(11): 3037-3044.
[40] BASHEER R, SHERIN JE, SAPER CB, et al.Effects of sleep on wake-induced c-fos expression [J]. J Neurosci, 1997, 17(24):9746-9750.
[41] SHERIN JE, SHIROMANI PJ, MCCARLEY RW, et al. Activation of ventrolateral preoptic neurons during sleep [J]. Science, 1996, 271(5246): 216– 219.
[42] PETERFI Z, CHURCHILL L, HAJDA I, et al A.Fos-immunoreactivity in the hypothalamus: dependency on the diurnal rhythm, sleep, gender, and estrogen [J]. Neuroscience, 2004, 124(3):695-707.
[43] SZYMUSIAK R, GVILIA I, MCGINTY D. Hypothalamic control of sleep [J]. Sleep Med, 2007, 8(4):291-301
[44] GVILIA I, ANGARA C, MCGINTY D, et al. Different neuronal populations of the rat median preoptic nucleus express c-fos during sleep and in response to hypertonic saline or angiotensin-II [J]. J Physiol, 2005, 569(Pt2):587-599.
[45] MODIRROUSTA M, MAINVILLE L, JONES BE. Gabaergic neurons with alpha2-adrenergic receptors in basal forebrain and preoptic area express c-fos during sleep [J]. Neuroscience, 2004, 129(3):803-810.
[46] GALLOPIN T, LUPPI PH, CAULI B, et al. The endogenous somnogen adenosine excites a subset of sleep-promoting neurons via A2A receptors in the ventrolateral preoptic nucleus [J]. Neuroscience, 2005, 134:1377–1390.
[47] KAUFLING J, VEINANTE P, PAWLOWSKI SA, et al. Afferents to the GABAergic tail of the ventral tegmental area in the rat [J]. J Comp Neurol, 2009, 513(6):597-621.
[48] FRONCZEK R, BAUMANN CR, LAMMERS GJ, et al. Hypocretin/ orexin disturbances in neurological disorders [J].Sleep Med Rev, 2009, 13 (1): 9-22.
[49] CHANG SY, KIM U. Ionic mechanism of long-lasting discharges of action potentials triggered by membrane hyperpolarization in the medial lateral habenula [J]. J Neurosci, 2004, 24(9): 2172-2181.
[2] PICKERING TG, DEVEREUX RB, JAMES GD, et al. Environmental infuences on blood pressure and the role of job train [J]. J Hypertens Suppl, 1996, 14 (5): 179-185.
[3] Sleep Disorders and Sleep Deprivation: an Unmet Public Health Problem. Institute of Medicine of the National Academies[C]. The National Academies Press, Washington DC, 2006.
[4] VON ECONOMO C. Sleep as a problem of localization [J]. J Nerv Ment Dis, 1930, 71: 249–259.
[5]姚泰.生理学(第五版)[M].北京:人民卫生出版社,2000:333.
[6] MONTI JM, JANTOS H. The roles of dopamine and serotonin, and of their receptors, in regulating sleep and waking [J]. Prog Brain Res, 2008, 172: 625-646.
[7] SZYMUSIAK R, McGINTY D. Hypothalamic regulation of sleep and arousal [J]. Ann N Y Acad Sci, 2008, 1129: 275-286.
[8] CESPUGLIO R, SARDA N, GHARIB A, et al. Voltammetric detection of the release of 5-hydroxyindole Compounds throughout the sleep-waking cycle of the rat [J]. Exp Brain Res, 1990, 80 (1): 121-128.
[9] PORTAS CM, BJORVATN B, FAGERLAND S. Detection of extracellular levels of serotonin in dorsal raphe nucleus and frontal cortex over the sleep/wake cycle in the freely moving rat [J]. Neuronscience, 1998, 83 (3): 807-814.
[10] JOUVET M. Sleep and neurotransmitters [J]. Rev Med Interne, 1996, 17: 18-23.
[11] STERIADE M, MCCARLEY RW. Brain control of sleep and wakefulness [M]. New York: Kluwer Academic Press, 2005.
[12] BERRIDGE CW, WATERHOUSE BD. The locus coeruleus-noradrenergic system: modulation of behavioral state and state-dependent cognitive processes. [J]. Brain Res Brain Res Rev, 2003, 42 (1): 33-84.
[13] THANKCHAN S, KAUR S, SHIROMANI PJ. Activity of pontine neurons during sleep and cataplexy in hypocretin knock-out mice. [J]. J Neurosci, 2009, 29 (5): 1580-1585.
[14] KOYAMA Y, TAKAHASHI K, KODAMA T, et al. State-dependent activity of neurons in the perifornical hypothalamic area during sleep and waking [J]. Neuroscience, 2003, 119 (4): 1209-1219.
[15] FRONCZEK R, BAUMANN CR, LAMMERS GJ, et al. Hypocretin/orexin disturbances in neurological disorders [J]. Sleep Medicine Reviews, 2009, 13 (1): 9-22.
[16] NAUTA WJH. Hypothalamic regulation of sleep in rats [J]. An experimentalstudy. J Neurophysiol, 1946, 9: 285–316.
[17] McGINTY D, STERMAN MB. Sleep suppression after basal forebrain lesions in the cat [J]. Science, 1968, 160: 1253–1255.
[18] SZYMUSIAK R, SATINOFF E. Ambient temperature-dependence of sleep disturbances produced by basal forebrain damage in rats [J]. Brain Res Bull, 1984, 12(3): 295–305.
[19] GONG H, SZYMUSIAK R, KING J, et al. Sleep-related c-Fos protein expression in the preoptic hypothalamus: effects of ambient warming [J]. Am J Physiol Regul Integr Comp Physiol, 2000, 279 (6): 2079-2088.
[20] SZYMUSIAK R, ALAM N, STEININGER TL, et al. Sleep-waking discharge patterns of ventrolateral preoptic/anterior hypothalamic neurons in rats [J]. Brain Res, 1998, 803 (1-2): 178–188.
[21] GONG H, McGINTY D, GUZMAN-MARIN R, et al. Activation of c-Fos in GABAergic neurones in the preoptic area during sleep and in response to sleep deprivation [J]. J Physiol, 2004, 556(Pt3): 935–946.
[22] PARAMENTIER R, KOLBAEV S, KLYUCH BP, et al. Excitation of histaminergic tuberomamillary neurons by thyrotropin-releasing hormone [J]. J Neurosci. 2009, 8, 29(14):4471-4483.
[23] MITANI A, ITO K, HALLANGER AE, et al. Cholinergic projections from the laterodorsal and pedunculopontine tegmental nuclei to the pontine gigantocellular tegmental field in the cat [J]. Brain Res, 1988, 451(1-2): 397–402.
[24] GNADT JW, PEGRAM GV. Cholinergic brainstem mechanisms of REM sleep in the rat [J]. Brain Res, 1986, 384 (1):29–41.
[25] STERIADE M, McCARLEY RW. Brain control of sleep and wakefulness.New York: Kluwer Academic Press, 2005.
[26] McCARLEY RW. Mechanisms and models of REM sleep control [J]. Archives Ital de Biol, 2004, 142: 429–468.
[27] FULLER PM, SAPER CB, LU J. The pontine REM switch: past and present [J]. J Physiol, 2007, 584(Pt 3): 735-741.
[28] GREENE RW, GERBER U, McCARLEY RW. Cholinergic activation of medial pontine reticular formation neurons in vitro [J]. Brain Res, 1989, 476: 154–159.
[29] FORNAL C, AUERBACH S, JACOBS BL. Activity of serotonin- containing neurons in nucleus raphe magnus in freely moving cats [J]. Exp Neurol, 1985,88(3): 590–608.
[30] LEVINE ES, JACOBS BL. Neurochemical afferents controlling theactivity of serotonergic neurons in the dorsal raphe nucleus: microiontophoretic studies in the awake cat [J]. J Neurosci, 1992, 12:4037–4044.
[31] GERVASONI D, DARRACQ L, FORT P, et al. Electrophysiological evidence that noradrenergic neurons of the rat locus coeruleus are tonically inhibited by GABA during sleep [J]. Eur J Neurosci, 1998, 10:964–970.
[32] JONES IS. The organization of central cholinergic systems and their functional importance in sleep–waking states[J]. Prog Brain Res, 1993, 98: 61–71.
[33] TORTEROLO P, MORALES FR, CHASE MH. GABAergic mechanisms in the pedunculopontine tegmental nucleus of the cat promote active (REM) sleep [J]. Brain Res, 2002, 944(1-2): 1-9.
[34] RADULOVACKI M. Role of adenosine in sleep in rats [J]. Rev Clin Basic Pharmacol, 1985, 5(3-4):327–339.
[35] TICHO SR, RADULOVACKI M. Role of AD in sleep and temperature regulation in the preoptic area of rats [J]. Pharmacol Biochem Behav, 1991, 40(1):33–40.
[36] PHILLIS JW, WU PH. The role of adenosine and its nucleotide in central synaptic transmission [J]. Prog Neurobiol, 1981, 16(3-4): 187–239.
[37] PHILLIS J.W., WALTER G.A., O’REGAN M.H., et al. Adenosine in the control of cerebral circulation [J]. Cerebrovasc. Brain Metab. Rev, 1989, 1: 26–54.
[38] TICHO SR, RADULOVACKI M. Role of adenosine in sleep and temperature regulation in the preoptic area of rats [J]. Pharmacol Biochem Behav, 1991, 40(1):33-40.
[39] FREDHOLM BB. Astra Award Lecture. Adenosine, adenosine receptors and the actions of caffeine [J]. Pharmacol Toxicol, 1995, 76(2):93-101.
[40] FREDHOLM BB, ARSLAN G, HALLDNER L, et al. Structure and function of adenosine receptors [J]. Naunyn Schmiedebergs Arch Pharmacol, 2000, 362(4-5): 364–374.
[41] KLOTZ KN. Adenosine receptors and their ligands [J]. Naunyn Schmiedebergs Arch Pharmacol, 2000, 362(4-5):382–391.
[42] PORKKA-HEISKANEN T, STRECKER RE, THAKKAR M, et al. Adenosine: a mediator of the sleep-inducing effects of prolonged wakefulness [J]. Science,1997, 276(5316): 1265– 1268.
[43] PORKKA-HEISKANEN T, STRECKER RE, McCARLEY RW. Brain site specificity of extracellular adenosine concentration changes during sleep deprivation and spontaneous sleep: an in vivo microdialysis study [J]. Neuroscience 2000, in press.
[44] RAINNIE DG, GRUNZE HC, McCARLEY RW, et al. Adenosine inhibition of mesopontine cholinergic neurons: implications for EEG arousal [J]. Science, 1994, 263:689–692.
[45] ROBERT E. STRECKER, STEPHEN MORAIRTY, MAHESH M THAKKER, et al .Adenosinergic modulation of basal forebrain and preoptic/anterior hypothalamic neuronal activity in the control of behavioral state[J]. Behavioural Brain Research, 2000, 115(2):183-204.
[46] MATSUMURA H, NAKAJIMA T, OSAKA T, et al. Prostaglandin D2-sensitive, sleep promoting zone defined in the ventral surface of the rostral basal forebrain [J]. Proc Natl Acad Sci USA, 1994, 91(25):11998–12002.
[47] SATOH S, MATSUMURA H, SUZUKI F, et al. Promotion of sleep mediated by the A2a-adenosine receptor and possible involvement of this receptor in the sleep induced by prostaglandin D2 in rats [J]. Proc Natl Acad Sci U S A., 1996, 93(12):5980-5984.
[48] MIZOGUCHI A, EGUCHI N, KIMURA K, et al. Dominant localization of prostaglandin D receptors on arachnoid trabecular cells in mouse basal forebrain and their involvement in the regulation of non-rapid eye movement sleep [J]. Proc Natl Acad Sci USA, 2001, 98(25):11674–11679.
[49] MORAIRTY S, RAINNIE D, McCARLEY R, et al. Disinhibition of ventrolateral preoptic area sleep-active neurons by adenosine: a new mechanism for sleep promotion [J]. Neuroscience, 2004, 123(2):451-457.
[50] ERIC E. ABRAHAMSON, ROBERT Y. MOORE. Suprachiasmatic nucleus in the mouse: retinal innervation, intrinsic organization and efferent projections [J]. Brain Research, 2001, 916(1-2): 172–191.
[51] LESAUTER J, LEHMAN MN, SILVER R. Restoration of circadian rhythmicity by transplants of SCN "micropunches"[J]. J Biol Rhythms, 1996, 11(2):163-171.
[52] IBUKA N, KAWAMURA H. Loss of circadian rhythm in sleepwakefulness cycle in the rat by suprachiasmatic nucleus lesions [J]. Brain Res, 1975, 96(1):76-81.
[53] MOORE RY, EICHLER VB. Loss of a circadian adrenal corticosterone rhythm following suprachiasmatic lesions in the rat [J]. Brain Res, 1972, 42(1):201-206.
[54]周先举,俞光第,印其章.哺乳动物昼夜节律组构中的下丘脑视交叉上核和松果腺.生理科学进展[J],2001,32(2):116-120.
[55] CHALLET E, CALDELAS I, GRAFF C, et al. Synchronization of the molecular clockwork by light- and food-related cues in mammals [J]. Biol Chem, 2003, 384(5):711-719.
[56] ZHAO H., RUSAK B. Circadian fring-rate rhythms and light responses of rat habenular nucleus neurons in vivo and in vitro [J]. Neuroscience, 2005, 132: 519–528.
[57] MATTHEWS-FELTON T, CORODIMAS KP, ROSENBLATT JS, et al. Lateral habenula neurons are necessary for the hormonal onset of maternal behavior and for the display of postpartum estrus in naturally parturient female rats [J]. Behav Neurosci, 1995, 109(6): 1172–1188.
[58] GOLDSTEIN RA. GABAergic habenulo-raphe pathway mediation of the hypnogenic effects of vasotocin in the cat [J]. Neuroscience, 1983, 10(3):941–945.
[59] GALLAGHER M., CHIBA AA. The amygdala and emotion [J]. Neurobiology, 1996, 6(2):221-227.
[60]王绍.缰核与奖赏和稳态[J].生理科学进展,2008,4(39): 292-296.
[61] HERKENHAM M., NAUTA W.J.H. Efferent connections of the habenular nuclei in the rat [J]. Journal of Comparative Neurology, 1979, 187(1): 19–47.
[62] ANDRES KH, VON DURING M, VEH RW. Subnuclear organization of the rat habenular complexes [J].The Journal of Comparative Neurology, 1999, 407(1): 130–150.
[63] KIM U, CHANG SY. Dendritic morphology, local circuitry and intrinsic electrophysiology of neurons in the rat medial and lateral habenular nuclei of the epithalamus [J]. J COMP NEURO, 2005, 483 (2): 236-250.
[64] ARAKI M, MCGEER PL, MCGEER, EG. Retrograde HRP tracing combined with a pharmacohistochemical method for GABA transaminase for the identification of presumptive GABAergic projections to the habenula [J]. Brain Research, 1984, 304(2): 271–277.
[65] GARLAND JC, MOGENSON GJ. An electrophysiological study of convergenceof entopeduncular and lateral preoptic inputs on lateral habenular neurons projecting to the midbrain [J]. Brain Research, 1983, 263: 33–41.
[66] VINCENT SR, KIMURA H, MCGEER EG. A histochemical study of GABA-transaminase in the efferents of the pallidum [J]. Brain Research, 1982, 241(1): 162–165.
[67] NAGY JI, CARTER DA, LEHMANN J, et al. Evidence for a GABA-containing projection from the entopedoncular nucleus to the lateral habenula in the rat [J]. Brain Research, 1978, 145(2):360–364.
[68] CHANG S, KIM U. Ionic mechanism of long-lasting discharges of action potentials triggered by membrane hyperpolarization in the medial lateral habenula [J]. Journal of Neuroscience, 2004, 24: 2172–2181.
[69] MORIIZUMI T, HATTORI T. Choline acetyltransferase- immunoreactive neurons in the rat entopedoncular nucleus [J]. Neuroscience, 1992, 46(3):721–728.
[70] VERTES RP, FORTIN WJ, CRANE AM. Projections of the median raphe′nucleus in the rat [J]. Journal of Comparative Neurology, 1999, 407: 555–582.
[71] QU T, DONG K, SUGIOKA K, et al. Demonstration of direct input from the retina to the lateral habenular nucleus in the albino rat [J]. Brain Research, 1996, 709(2): 251–258.
[72] KAWAJA MD, FLUMERFEIL BA., HRYCYSHYN AM. Synaptic organization of septal projections in the rat medial habenula: a wheat germ agglutinin-horseradish peroxidase and immunohistochemical study [J]. Synapse, 1990, 6(1): 45–54.
[73] HERKENHAM M, NAUTA WJH. Afferent connections of the habenular nuclei in the rat. A horseradish peroxidase study, with a note on the fiber-of-passage problem [J]. Journal of Comparative Neurology, 1977, 173(1): 123–146.
[74] LECOURTIER L, KELLY PH. A conductor hidden in the orchestra? Role of the habenular complex inmonoamine transmission and cognition [J]. Neurosci Biobehav Rev, 2007, 31(5):658-672.
[75] HIKOSAKA O, SESACK SR, LECOURTIER L, et al. Habenula: crossroad between the basal ganglia and the limbic system [J]. J Neurosci, 2008, 28(46):11825-11829.
[76] GOLDSTEIN R. Vasotocin-like activity of cerebrospinal fluid induced byhabenular lesions in cats [J]. Neuropeptides, 1985, 6(4):303-310.
[77] MATTHEWS-FELTON T, LINTON L, ROSENBLATT JS, et al. First and second order maternal behavior related afferents of the lateral habenula [J]. Neuroreport, 1999, 10(4):883–887.
[78] SZYMUSIAK R, STEININGER T, ALAM N, et al. Preoptic area sleep-regulating mechanisms [J]. Arch Ital Biol, 2001, 139(1-2):77-92.
[79] FERRARO G, MONTALBANO ME, SARDO P, et al. Lateral habenula and hippocampus: a complex interaction raphe cells mediated [J]. Journal of Neural Transmission, 1997, 104(6-7): 615–631.
[80] ULLSPERGERM, YVES VON CRAMON D. The error monitoring using external feedback: specific roles of the habenular complex, the reward system, and the cingulate mot or area revealed by functi onal magnetic resonance imaging [J]. J Neurosci, 2003, 23: 4308-4314.
[81] JI H, SHEPARD PD. Lateral Habenula Stimulati on Inhibits Rat Midbrain Dopamine Neurons through a GABAA Receptor Mediated Mechanism [J]. J Neurosci, 2007, 27: 6923～6930 .
[82] MATSUMOTO M, HIKOSAKA O. Lateral habenula as a source of negative reward signals in dopamine neurons [J]. Nature, 2007, 447(7148): 1111-1115.
[83]赵华,王绍.分别兴奋大鼠内侧和外侧缰核引起痛阈的不同改变[J].生理学报,1995, 3(47): 292-296.
[84] YANG LM, HU B, XIA YH, et al. Lateral habenula lesions improve the behavioral response in depressed rats via increasing the serotonin level in dorsal raphe nucleus [J]. Behav Brain Res, 2008, 188(1):84-90.
[85] HAUN F, ECKENRODE TC, MURRAY M. Habenula and thalamus cell transplants restore normal sleep behaviors disrupted by denervation of the interpeduncular nucleus [J]. J Neurosci, 1992, 12(8):32.
[86] KELEIN DC, WELLER JL. Rapid light-induced decrease in pineal seroton N-acetyltransferase activity [J].Science, 1972, 177(48):532-533.
[87] LIU C, WEAVER DR, JIN X, et al. Molecular dissection of two distinct actions of melatonin on thesuprachiasmatic circadian clock [J].Neuron, 1997, 19:91-102.
[88] KOKKOLA T, LAITINEN JT. Melatonin receptor genes [J]. Ann Med, 1998, 30(1): 88-94.
[89]吴建群,王绍.缰核与GABA、乙酰胆碱[J].针刺研究,1989,1-2: 72- 73.
[90] CLAUDE GOTTESMANN. GABA mechanisms and sleep [J]. Neuroscience, 2002, 111(2): 231- 239.
[1] YANG LM, HU B, XIA YH, et al. Lateral habenula lesions improve the behavioral response in depressed rats via increasing the serotonin level in dorsal raphe nucleus [J]. Behav Brain Res, 2008, 188(1): 84-90.
[2] MORISSETTE MC, BOYE SM. Electrolytic lesions of the habenula attenuate brain stimulation reward [J]. Behav Brain Res, 2008, 187(1): 17-26.
[3] SARTORIUS A, HENN FA. Deep brain stimulation of the lateral habenula intreatment resistant major depression [J]. Med Hypotheses, 2007, 69(6): 1305-1308.
[4] ZHAO H, RUSAK B. Circadian firing-rate rhythms and light responses of rat habenular nucleus neurons in vivo and in vitro [J]. Neuroscience, 2005, 132(2): 519-528.
[5] KIM U, CHANG SY. Dendritic morphology, local circuitry and intrinsic electrophysiology of neurons in the rat medial and lateral habenular nuclei of the epithalamus [J]. J Comp Neurol, 2005, 483(2): 236-250.
[6] GOLDSTEIN R.A. GABAergic habenulo-raphe pathway mediation of the hypnogenic effects of vasotocin in cat [J]. Neuroscience, 1983, 10(3): 941-945.
[7] FLUMERFELT BA, CONTESTABILE A. Acetylcholinesterase Histochemistry of the habenulo-interpeduncular pathway in the rat and the effects of electrolytic and kainic acid lesions [J]. Anat Embryol (Berl), 1982, 163(4):435-446.
[8] IWAKIRI H, MATSUYAMA K, MORI S. Extracellular levels of serotonin in the medial pontine reticular formation in relation to sleep-wake cycle in cats: a microdialysis study [J]. Neurosci Res, 1993, 18(2):157-170.
[9] KOWSKI AB, GEISLER S, KRAUSS M, et al. Differential projections from subfields in the lateral preoptic area to the lateral habenular complex of the rat[J].J Comp Neurol, 2008, 507(4):1465-1478.
[10] ANDRES KH, VON DüRING M, VEH RW. Subnuclear organization of the rat habenular complexes [J]. J Comp Neurol, 1999, 407(1):130-150.
[11] GEISLER S, ANDRES KH, VEH RW. Morphologic and cytochemical criteria for the identification and delineation of individual subnuclei within the lateral habenular complex of the rat [J]. J Comp Neurol, 2003, 458(1):78-97.
[12] STAINES WA, YAMAMOTO T, DEWAR KM, et al. Distribution, morphology and habenular projections of adenosine deaminase-containing neurons in the septal area of rat [J]. Brain Res, 1988, 455(1):72-87.
[13] MATTEAU I, BOIRE D, PTITO M. Retinal projections in the cat: a cholera toxin B subunit study [J].Vis Neurosci, 2003, 20(5):481-493.
[14] SHIN TOKUNAGA, YASUHIRO TAKEDA, KAZUAKI SHINAMIYA, et al. Effects of some H1-antagonists on the sleep-Wake cycle in sleep-disturbed rats [J]. J Pharmacol Sci, 2007, 103(2):201-206.
[15] SHINOMIYA K, INOUE T, UTSU Y, et al. Effects of kava-kava extract on thesleep-wake cycle in sleep-disturbed rats [J]. Psychopharmacology (Berl), 2005, 180(3): 564-569.
[16] NECKELMANN D, URSIN R. Sleep stages and EEG power spectrum in relation to acoustical stimulus arousal threshold in the rat [J]. Sleep, 1993, 16(5): 467-477.
[17] FAULHABER J, STAIGER A, LANCEL M. The GABAA agonist THIP produces slow wave sleep and reduces spindling activity in NREM sleep in humans [J]. Psychopharmacology (Berl), 1997, 130(3):285–291.
[18] LANCEL M. The GABA (A) agonist THIP increases non-REM sleep and enhances non-REM sleep-specific delta activity in the rat during the dark period. Sleep, 1997, 20(12):1099–1104.
[19] ELLISON G. Stimulant-induced psychosis, the dopamine theory of schizophrenia, zand the habenula [J]. Brain Res Brian Res Rev, 1994, 19(2):223-239.
[20] HUSTON JP, HAAS HL, BOIX F, et al. Extracellular adenosine levels in neostriatum and hippocampusduring rest and activity periods of rats [J]. Neuroscience, 1996, 73(1):99-107.
[21] PORKKA-HEISKANEN T, STRECKER RE, MCCARLEY RW. Brain site-specificity of extracellular adenosine concentration changes during sleep deprivation and spontaneous sleep: an in vivo microdialysis study [J]. Neuroscience, 2000, 99(3):507-517.
[22] STRECKER RE, MORAIRTY S, THAKKAR MM, et al. Adenosinergic modulation of basal forebrain and preoptic/anterior hypothalamic neuronal activity in the control of behavioral state[J]. Behav Brain Res, 2000, 115(2):183-204.
[23] MURILLO-RODRIGUEZ E, BLANCO-CENTURION C, GERASHCHENKO D, et al. The diurnal rhythm of adenosine levels in the basal forebrain of young and old rats [J]. Neuroscience, 2004, 123(2):361-370.
[24] PORTAS CM, THAKKAR M, RAINNIE DG, et al. Role of adenosine in behavioral state modulation: a microdialysis study in the freely moving cat [J]. Neuroscience, 1997, 79(1):225-235.
[25] LUCAS EA, STERMAN MB. Effects of a forebrain lesion on the poly-cyclic sleep–wake cycle and sleep–wake patterns in the cat [J]. Exp Neurol, 1975, 46: 368–388.
[26] SZYMUSIAK R, McGINTY D. Sleep suppression following kainic acid-induced lesions of the basal forebrain [J]. Exp Neurol, 1986, 94:598–614.
[27] SALLANON M, DENOYER M, KITAHAMA K, et al. Long lasting insomnia induced by preoptic neuron lesions and its tran-sient reversal by muscimol injection into the posterior hypothalamus in the cat [J]. Neuroscience, 1989, 32: 669–683.
[28] SRIVIDYA R, MALLICK HN, KUMAR VM. Differences in the effects of medial and lateral preoptic lesions on thermoregulation and sleep in rats [J]. Neuroscience, 2006, 139(3):853-864.
[29] LECOURTIER L, NEIJT HC, KELLY PH. Habenula lesions cause impaired cognitive performance in rats: implications for schizophrenia [J]. Eur J Neurosci, 2004, 19(9):2551–2560.
[30] SHEPARD PD, HOLCOMB HH, GOLD JM. The presence of absence: Habenular regulation of dopamine neurons and the encoding of negative outcomes [J]. Schizophrenia Bull, 2006, 32(3):417–421.
[31] SHUMAKE J, EDWARDS E, GONZALEZ-LIMA F. Opposite metabolic changes in the habenula and ventral tegmental area of a genetic model of helpless behavior [J]. Brain Res, 2003, 963:274–281.
[32] FELTON T, LINTON L, ROSENBLATT JS, et al. First and second order maternal behavior related afferents of the lateral habenula [J]. Neuroreport, 1999, 10(4):883-887.
[33] BRISKI K, GILLEN E. Differential distribution of Fos expression within the male rat preoptic area and hypothalamus in response to physical vs. psychological stress [J]. Brain Res Bull, 2001, 55(3): 401-408.
[34] GOLDSTEIN R. Vasotocin-like activity of cerebrospinal fluid induced by habenular lesions in cats [J]. Neuropeptides, 1985, 6(4):303-310.
[35] GARLAND JC, MOGENSON GJ. An Electrophysiological Study of Convergence of Entopeduncular and LateralPreoptic Inputs on Lateral Habenular Neurons Projecting to the Midbrain [J]. Brain Research, 1983, 263(1):33-41.
[36] METHIPPARA MM, KUMAR S, ALAM MN, et al. Effects on sleep of microdialysis of adenosine A1 and A2A receptor analogs into the lateral preoptic area of rats [J]. Am J Physiol Regul Integr Comp Physiol, 2005, 89(6): 1715–1723.
[37] GVILIA I, FENG XU, MCGINTY D et al. Homeostatic Regulation of Sleep: A Role for Preoptic Area Neurons [J]. The Journal of Neuroscience, 2006, 26(37): 9426–9433.
[38] DéTáRI L, JUHáSZ G, KUKORELLI T. Firing properties of cat basal forebrain neurones during sleep-wakefulness cycle [J]. Electroencephalogr Clin Neurophysiol, 1984, 58(4):362-368.
[39] GVILIA I, TURNER A, MCGINTY D, et al. Preoptic area neurons and the homeostatic regulation of rapid eye movement sleep [J]. J Neurosci, 2006, 26(11): 3037-3044.
[40] BASHEER R, SHERIN JE, SAPER CB, et al.Effects of sleep on wake-induced c-fos expression [J]. J Neurosci, 1997, 17(24):9746-9750.
[41] SHERIN JE, SHIROMANI PJ, MCCARLEY RW, et al. Activation of ventrolateral preoptic neurons during sleep [J]. Science, 1996, 271(5246): 216– 219.
[42] PETERFI Z, CHURCHILL L, HAJDA I, et al A.Fos-immunoreactivity in the hypothalamus: dependency on the diurnal rhythm, sleep, gender, and estrogen [J]. Neuroscience, 2004, 124(3):695-707.
[43] SZYMUSIAK R, GVILIA I, MCGINTY D. Hypothalamic control of sleep [J]. Sleep Med, 2007, 8(4):291-301
[44] GVILIA I, ANGARA C, MCGINTY D, et al. Different neuronal populations of the rat median preoptic nucleus express c-fos during sleep and in response to hypertonic saline or angiotensin-II [J]. J Physiol, 2005, 569(Pt2):587-599.
[45] MODIRROUSTA M, MAINVILLE L, JONES BE. Gabaergic neurons with alpha2-adrenergic receptors in basal forebrain and preoptic area express c-fos during sleep [J]. Neuroscience, 2004, 129(3):803-810.
[46] GALLOPIN T, LUPPI PH, CAULI B, et al. The endogenous somnogen adenosine excites a subset of sleep-promoting neurons via A2A receptors in the ventrolateral preoptic nucleus [J]. Neuroscience, 2005, 134:1377–1390.
[47] KAUFLING J, VEINANTE P, PAWLOWSKI SA, et al. Afferents to the GABAergic tail of the ventral tegmental area in the rat [J]. J Comp Neurol, 2009, 513(6):597-621.
[48] FRONCZEK R, BAUMANN CR, LAMMERS GJ, et al. Hypocretin/ orexin disturbances in neurological disorders [J].Sleep Med Rev, 2009, 13 (1): 9-22.
[49] CHANG SY, KIM U. Ionic mechanism of long-lasting discharges of action potentials triggered by membrane hyperpolarization in the medial lateral habenula [J]. J Neurosci, 2004, 24(9): 2172-2181.