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mise à jour du 30 mai 2002
Behavioural Brain Research
2002;28:169-177
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Cortical arousal induced by microinjection of orexins into the paraventricular nucleus of the rat
Ikuko Sato-Suzuki, Ichiro Kita, Yoshinari Seki,
Mitsugu Oguri, Hideho Arita
Department of Physiology, Toho University School of Medicine, Tokyo, Japan
 
Voir chapitre sur l'hypocrétine et schéma de l'hypothalamus
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-Kita I, Kubota N, Yanagita S, Motoki C Intracerebroventricular administration of corticotropin-releasing factor antagonist attenuates arousal response accompanied by yawning behavior in rats. Neurosci.Letter 2008;433(3):205-208 
-Kita I, Yoshida Y, Nishino S. An activation of parvocellular oxytocinergic neurons in the paraventricular nucleus in oxytocin-induced yawning and penile erection. Neurosci Res. 2006;54(4):269-275
-Kita I, Seki Y, Nakatani Y, Fumoto M, Oguri M, Sato-Suzuki I, Arita H. Corticotropin-releasing factor neurons in the hypothalamic paraventricular nucleus are involved in arousal/yawning response of rats. Behav Brain Res. 2006;169(1)48-56.
-Kita I, Sato-Suzuki et al.Yawning responses induced by local hypoxia in the paraventricular nucleus of the rat.Behavioural Brain Research 2000;117(1-2):119-126
-Kubota N, Amemiya S, Motoki C, Otsuka T, Nishijima T, Kita I. Corticotropin-releasing factor antagonist reduces activation of noradrenalin and serotonin neurons in the locus coeruleus and dorsal raphe in the arousal response accompanied by yawning behavior in rats. Neurosci Res. 2012;72(4):316-323
-Kubota N, Amemiya S, Yanagita S, Nishijima T, Kita I. Emotional stress evoked by classical fear conditioning induces yawning behavior in rats. Neurosci Lett. 2014 Mar 11.
-Seki Y, Y Nakatani, et al Light induces cortical activation and yawning in rat Behav Brain Res 2003;140(1-2):65-73
-Seki Y, Sato-Suzuki I, et al Yawning/cortical activation induced by microinjection of histamine into the paraventricular nucleus of the rat. Behav Brain Res. 2002;134(1-2):75-82.
-Sato-Suzuki I, Kita I, Oguri M, Arita H Stereotyped yawning responses induced by electrical and chemical stimulation of paraventricular nucleus of the rat Journal of Neurophysiology, 1998;80(5)2765-2775
-Sato-Suzuki I, I Kita, Seki Y, M Oguri, H Arita Cortical arousal induced by microinjection of orexins into the paraventricular nucleus of the rat Behavioural Brain Research 2002;128:169-177

Introduction : We recently reported that a stereotyped yawning response can be evoked by microinjection of L-glutamate, cyanide and a nitric oxide-releasing compound (NO) into the medial parvocellular subdivision of the paraventricular nucleus (PVN) in anesthetized, spontaneously breathing rats. In those studies, we recorded the electrocorticogram (ECoG) to evaluate arousal responses during yawning, and found that ECoG arousal, represented by lower voltage and faster rhythms, occurred before the final yawning behavior during both spontaneous yawning and yawning responses evoked by chemical stimulation. This arousal effect was of considerable significance because the data were obtained under anesthesia. Although the PVN is essential for the occurrence of yawning, our previous studies raised the possibility that the PVN may play an important role in triggering arousal mechanisms. We therefore suggested that the yawning model is appropriate for investigating arousal mechanisms in rats.
 
Orexins (A and B) are a recently identified family of neuropeptides originally believed to be important mediators of food intake. Apart from appetite regulation, orexins have now emerged as important regulators of sleep and arousal. For example, Chernelli et al. proposed that orexin knockout mice may be a model for human narcolepsy. They further observed that orexin neurons innervate neurons critical for controlling arousal in the ascending cortical activation system. Hagan et al. demonstrated that orexin-A activates locus coeruleus cell firing in vitro and increases arousal in rats in vivo. Moreover, Piper et al. reported that orexin-A modifies the sleep wake cycle of rats. In those studies, orexin-A stimulated arousal after intracerebroventricular injection, but the precise site of action within the brain for orexin-A to stimulate arousal remains unknown. We sought to determine whether the PVN is orexin sensitive by microinjecting orexin-A into the specific region within the PVN where a stereotyped yawning response was induced by L-glutamate. The effect of orexin-B microinjection into the PVN was also examined. The doses of orexin-A and orexin-B used in the present study were at least 1000 times smaller than those used for feeding responses in microinjection studies of the PVN. All microinjections were done with multi-barrel glass micropipettes which provide especially high resolution, thus stimulating only a restricted region within the PVN.
orexin
[....] The yawning response has important physiological significance for arousal in anesthetized animals. It is apparent from observing human subjects that yawning is not a behavior restricted to mouth opening, but is a coordinated motor pattern characterized by a deep inspiration and stretching of the trunk. Yawning is also accompanied by changes in autonomic function, including a depressor response, lacrimation and erection. Furthermore, yawning is a phenomenon that subserves arousal.
 
To evaluate these various physiological aspects of yawning, we monitored polygraphic measures representing a yawning response in anesthetized, spontaneously breathing rats. It is important to emphasize that if we used conscious animals, the data obtained would be almost certainly restricted to counting the number of mouth openings, which shows only one aspect of the entire yawning response. Whereas the ECoG of anesthetized rats is normally characterized by high voltage and very slow waves (2Hz), faster rhythms (4Ð6 Hz) became dominant afterchemical stimulation of the PVN.
 
Since this arousal shift in the ECoG was usually accompanied by yawning responses, we suggest that the PVN mediates the cortical activation related to yawning. In this respect, Dringenberg and Vanderwolf demonstrated that several pathways contribute to ECoG activation as well as arousal mechanisms. Our findings indicate that the PVN plays a significant role in the arousal mechanism related to yawning. Narcolepsy, a disorder characterized by excessive daytime sleepiness, cataplexy, and striking transitions from wakefulness into rapid eye movement sleep, is apparently caused by orexin and orexin receptor deficiency in human and animal models. The type of orexin receptor deficiency in this disorder is OX2R, which is predominantly found in the PVN. These data together suggest orexin as a major neurotransmitter involved in sleep and arousal mechanisms, and further strengthen our hypothesis that the PVN plays an important role in arousal systems.
 
In conclusion, microinjection of orexin-A into the PVN elicited an arousal/yawning response, whereas orexin-B injection induced the arousal response alone. The results demonstrate that an orexin receptive site for triggering arousal/yawning responses exists in the PVN. Our finding should open a new avenue of research to investigate the role of the PVN in arousal regulation.
 
-Kita I, Kubota N, Yanagita S, Motoki C Intracerebroventricular administration of corticotropin-releasing factor antagonist attenuates arousal response accompanied by yawning behavior in rats. Neurosci. Lettre 2008; 
-Kita I, Yoshida Y, Nishino S. An activation of parvocellular oxytocinergic neurons in the paraventricular nucleus in oxytocin-induced yawning and penile erection. Neurosci Res. 2006;54(4):269-275
-Kita I, Seki Y, Nakatani Y, Fumoto M, Oguri M, Sato-Suzuki I, Arita H. Corticotropin-releasing factor neurons in the hypothalamic paraventricular nucleus are involved in arousal/yawning response of rats. Behav Brain Res. 2006; 169; 1; 48-56.
-Kita I, Sato-Suzuki et al.Yawning responses induced by local hypoxia in the paraventricular nucleus of the rat.Beh Brain Res 2000; 117; 1-2; 119 - 126
-Sato-Suzuki I, Kita I; Oguri M, Arita H Stereotyped yawning responses induced by electrical and chemical stimulation of paraventricular nucleus of the rat Journal of Neurophysiology, 1998; 80, 5; 2765-2775
-Sato-Suzuki I, I Kita, YSeki, M Oguri, H Arita Cortical arousal induced by microinjection of orexins into the paraventricular nucleus of the rat Behav Brain Res 2002; 128; 169-177
-Seki Y, Y Nakatani, et al Light induces cortical activation and yawning in rat Behav Brain Res 2003; 140; 1-2; 65-73
-Seki Y, Sato-Suzuki I, et al Yawning/cortical activation induced by microinjection of histamine into the paraventricular nucleus of the rat. Behav Brain Res. 2002;134(1-2):75-82.
-Collins G, JM Witkin et al Dopamine agonist-induced yawning in rats: a dopamine d3 receptor mediated behavior J Pharmacol Exp Ther 2005
-Hipolide DC; Lobo LL; De Medeiros R; Neumann B; Tufik S Treatment with dexamethasone alters yawning behavior induced by cholinergic but not dopaminergic agonist. Physiol Behav 1999; 65; 4-5; 829-32
-Hipolide DC, Tufik S Paradoxical sleep deprivation in female rats alters drug-induced behaviors Physiol Behav. 1995; 57; 6; 1139-1143
-Moyaho A, Valencia J Grooming and yawning trace adjustment to unfamiliar environments in laboratory Sprague-Dawley rats J Comp Psychol 2002; 116; 3; 263-269
-Neumann BG, Troncone LR, Braz S, Tufik S Modifications on dopaminergic and cholinergic systems induced by the water tank technique: analysis through yawning behavior. Arch Int Pharmacodyn Ther 1990; 308; 32-38
-Tufik S et al Effects of stress on drug induced yawning Physiol Behav 1995; 58; 1; 1881-1884
 
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