229:
tubermammillary nucleus, raphe nuclei, and locus coeruleus. Thus, they are inhibited by histamine, serotonin, and norepinepherine. This mutual inhibition may provide the basis for establishing periods of sleep and waking. A reciprocal inhibition also characterizes an electronic circuit known as the flip-flop. A flip-flop can assume one of two states, usually referred to as on or off. Thus, either the sleep neurons are active and inhibit the wakefulness neurons, or the wakefulness neurons are active and inhibit the sleep neurons, Because these regions are mutually inhibitory, it is impossible for neurons in both sets of regions to be active at the same time. This flip-flop, switching from one state to another quickly, can be unstable.
209:, two known inhibitory neurotransmitters, to arousal-promoting neurons, such as histaminergic, serotonergic, orexinergic, noradrenergic, and cholinergic neurons (neurons mentioned above). Levels of acetylcholine, norepinephrine, serotonin, and histamine decrease with the onset of sleep, for they are all wakefulness promoting neurotransmitters. Therefore, it is believed that the activation of sleep-promoting neurons causes the inhibition of arousal-promoting neurons, which leads to sleep. Evidence has shown that during the sleep-wake cycle, sleep-promoting neurons and the arousal-promoting neurons have reciprocal discharges, and that during
228:
The majority of sleep neurons are located in the ventrolateral preoptic area (vlPOA). These sleep neurons are silent until an individual shows a transition from waking to sleep. The sleep neurons in the preoptic area receive inhibitory inputs from some of the same regions they inhibit, including the
76:
Von
Economo, in his studies, noticed that lesions in the connection between the midbrain and the diencephalon caused prolonged sleepiness and therefore proposed the idea of an ascending arousal system. During the past few decades major ascending pathways have been discovered with located neurons and
237:
The sleep cycle is normally defined in stages. When an individual first begins to sleep, stage 1 is entered, marked by the presence of some theta activity, which indicates that the firing of neurons in the neocortex is becoming more synchronized, as well as alpha wave activity (smooth electrical
224:
Adenosine is also known as the sleep promoting nucleoside neuromodulator. Astrocytes maintain a small stock of nutrients in the form of glycogen. In times of increased brain activity, such as during daytime, this glycogen is converted into fuel for neurons; thus, prolonged wakefulness causes a
67:
Sleep electrophysiological measurements can be made by attaching electrodes to the scalp to measure the electroencephalogram (EEG) and to the chin to monitor muscle activity, recorded as the electromyogram (EMG). Electrodes attached around the eyes monitor eye movements, recorded as the
50:
decided to study this disease and identified a key component in the sleep-wake regulation. He identified the pathways that regulated wakefulness and sleep onset by studying the parts of the brain that were affected by the disease and the consequences it had on the
225:
decrease in the level of glycogen in the brain. A fall in the level of glycogen causes an increase in the level of extracellular adenosine, which has an inhibitory effect in neural activity. This accumulation of adenosine serves as a sleep-promoting substance.
429:
Gallopin, T; Luppi, PH; Cauli, B; Urade, Y; Rossier, J; Hayaishi, O; Lambolez, B; Fort, P (2005). "The endogenous somnogen adenosine excites a subset of sleep-promoting neurons via A2A receptors in the ventrolateral preoptic nucleus".
465:
Sallanon M, Denoyer M, Kitahama K, Aubert C, Gay N, Jouvet M (1989). "Long-lasting insomnia induced by preoptic neuron lesions and its transient reversal by muscimol injection into the posterior hypothalamus in the cat".
238:
activity of 8–12 Hz recorded from the brain, generally associated with a state of relaxation). This stage is a transition between sleep and wakefulness. This stage is classified as non-REM sleep.
46:
and attacked the part of the brain that regulates sleep influenced Europe and North
America. Although the virus that caused this disorder was never identified, the psychiatrist and neurologist
59:. His discoveries were not appreciated until the last two decades of the 20th century when the pathways of sleep were found to reside in the exact place that Constantin von Economo stated.
81:. This pathway divides into two branches: one that ascends to the thalamus and activates the thalamus relay neurons, and another one that activates neurons in the lateral part of the
109:. These neurons play a crucial role in bridging information in between the thalamus and the cerebral cortex. These neurons have high activation during wakefulness and during
145:
produce serotonin, dopamine and histamine respectively. They then project onto the hypothalamic peptidergic neurons, which contain melanin-concentrated hormones or
501:
Swett CP, Hobson JA (September 1968). "The effects of posterior hypothalamic lesions on behavioral and electrographic manifestations of sleep and waking in cats".
1401:
1019:
363:
Zhi-Li Haung; Yoshihiro Urade; Osamu
Hayaishi (2009). "Prostanglan and adenosine in the regulation of sleep and wakefulness".
344:
181:
while those in the posterior hypothalamus lead to sleepiness. Further research has shown that the hypothalamic region called
981:
565:"Characterization and mapping of sleep-waking specific neurons in the basal forebrain and preoptic hypothalamus in mice"
859:
854:
826:
528:
Saper CB, Chou TC, Scammell TE (December 2001). "The sleep switch: hypothalamic control of sleep and wakefulness".
1014:
971:
945:
704:
161:. It has also been discovered that lesions to this part of the brain cause prolonged sleep or may produce coma.
198:
182:
106:
1286:
1094:
956:
966:
894:
739:
651:
Saper, CB; TC Chou; TE Scammell (2001). "The sleep twitch: Hypothalamic control of sleep and wakefulness".
23:
1099:
961:
217:
receptors increase in the arousal-promoting neurons. This had led some to believe that the increase of
277:
Saper, C.B.; Scammell, T.E.; Lu, J. (2005). "Hypothalamic regulation of sleep and circadian rhythms".
1177:
986:
934:
909:
55:. He stated that the pathways that regulated sleep onset are located between the brain stem and the
734:
336:
142:
126:
31:
1274:
1055:
864:
1370:
919:
754:
47:
117:. The second branch originates from monoaminorgenic neurons. These neurons are located in the
1192:
924:
328:
169:
Some light was thrown on the mechanisms on sleep onset by the discovery that lesions in the
1346:
286:
8:
1341:
1331:
797:
697:
329:
247:
94:
290:
1351:
1104:
1004:
869:
628:
619:
603:
580:
443:
664:
541:
1361:
668:
633:
584:
545:
510:
483:
479:
447:
408:
340:
302:
78:
564:
1356:
1296:
660:
623:
615:
576:
537:
475:
439:
398:
294:
102:
52:
1227:
1154:
1132:
1079:
839:
744:
221:
receptors in the arousal-promoting neurons is another pathway of inducing sleep.
158:
130:
118:
90:
86:
56:
22:
is the transition from wakefulness into sleep. Sleep onset usually transits into
1396:
1301:
1172:
1114:
817:
717:
690:
1390:
1279:
1127:
1084:
1034:
1029:
802:
170:
154:
134:
98:
682:
1247:
1024:
1009:
672:
637:
588:
549:
451:
412:
306:
174:
138:
122:
82:
43:
514:
487:
1306:
1182:
1122:
1074:
1060:
899:
849:
844:
726:
362:
298:
1365:
1311:
1291:
1089:
1065:
996:
914:
807:
777:
772:
762:
252:
210:
129:. Each group produces a different neurotransmitter. The neurons in the
114:
27:
1375:
1326:
1321:
1187:
1149:
886:
792:
782:
767:
110:
428:
1242:
1217:
1137:
929:
904:
178:
197:
Sleep onset is induced by sleep-promoting neurons, located in the
1269:
1264:
1259:
1254:
1237:
1142:
976:
876:
834:
787:
403:
386:
206:
1336:
1222:
276:
146:
93:. This refers to the ascending reticular activating system (cf
1232:
1050:
713:
464:
201:(VLPO). The sleep-promoting neurons are believed to project
358:
356:
218:
214:
202:
186:
150:
30:) it is possible to transit from wakefulness directly into
335:. University of Massachusetts, Amherst: Pearson. pp.
1316:
1212:
601:
353:
272:
270:
268:
650:
380:
378:
562:
384:
97:). The cell group involved in the first pathway is an
424:
422:
265:
189:
that inhibits the arousal system during sleep onset.
375:
26:(NREM sleep) but under certain circumstances (e.g.
419:
322:
320:
318:
316:
604:"Afferents to the Ventrolateral Preoptic Nucleus"
527:
42:During the 1920s an obscure disorder that caused
1388:
563:Takahashi, K; JS Lin; K Sakai (June 16, 2009).
313:
137:, as fore the neurons in the dorsal and median
387:"Hypothalamic regulation of sleep and arousal"
712:
698:
602:Chou, TC; AA Bjorkum; SE Gaus; J Lu (2002).
149:, and basal forebrain neurons which contain
500:
1020:Rapid eye movement sleep behavior disorder
705:
691:
458:
141:, ventral periaqueductal grey matter, and
125:, ventral periaqueductal grey matter, and
627:
402:
185:produces the inhibitory neurotransmitter
326:
1389:
157:. These neurons then project onto the
16:Transition from wakefulness into sleep
686:
521:
192:
13:
620:10.1523/JNEUROSCI.22-03-00977.2002
581:10.1016/j.neuroscience.2009.02.075
444:10.1016/j.neuroscience.2005.05.045
14:
1413:
1402:Unsolved problems in neuroscience
385:McGinty, D; Szymusiak, R (2008).
62:
860:Obesity hypoventilation syndrome
855:Central hypoventilation syndrome
1015:Periodic limb movement disorder
982:Non-24-hour sleep–wake disorder
644:
595:
503:Archives Italiennes de Biologie
556:
494:
199:ventrolateral preoptic nucleus
183:ventrolateral preoptic nucleus
107:laterodorsal tegmental nucleus
1:
1287:Biphasic and polyphasic sleep
1095:Nocturnal clitoral tumescence
957:Advanced sleep phase disorder
665:10.1016/s0166-2236(00)02002-6
542:10.1016/S0166-2236(00)02002-6
258:
101:-producing cell group called
967:Delayed sleep phase disorder
895:Excessive daytime sleepiness
480:10.1016/0306-4522(89)90289-3
113:and a low activation during
24:non-rapid eye movement sleep
7:
1100:Nocturnal penile tumescence
972:Irregular sleep–wake rhythm
608:The Journal of Neuroscience
241:
71:
10:
1418:
962:Cyclic alternating pattern
232:
164:
37:
1205:
1178:Behavioral sleep medicine
1165:
1113:
1043:
995:
987:Shift work sleep disorder
943:
935:Sleep state misperception
885:
825:
816:
753:
724:
68:electro-oculogram (EOG).
735:Rapid eye movement (REM)
143:tuberomammillary nucleus
127:tuberomammillary nucleus
32:rapid eye movement sleep
1056:Exploding head syndrome
865:Obstructive sleep apnea
653:Trends in Neurosciences
530:Trends in Neurosciences
391:Frontiers in Bioscience
1371:Sleeping while on duty
920:Idiopathic hypersomnia
331:Physiology of Behavior
327:Carlson, Neil (2013).
48:Constantin von Economo
1193:Neuroscience of sleep
925:Night eating syndrome
910:Kleine–Levin syndrome
365:Journal of Physiology
89:, and throughout the
1347:Sleep and creativity
121:, dorsal and median
1342:Sleep and breathing
798:Sensorimotor rhythm
299:10.1038/nature04284
291:2005Natur.437.1257S
285:(7063): 1257–1264.
248:Sleep onset latency
95:reticular formation
1352:Sleep and learning
1105:Nocturnal emission
1005:Nightmare disorder
870:Periodic breathing
1384:
1383:
1362:Sleep deprivation
1201:
1200:
346:978-0-205-23939-9
79:neurotransmitters
1409:
1357:Sleep and memory
1297:Circadian rhythm
1044:Benign phenomena
946:Circadian rhythm
823:
822:
707:
700:
693:
684:
683:
677:
676:
648:
642:
641:
631:
599:
593:
592:
560:
554:
553:
525:
519:
518:
498:
492:
491:
462:
456:
455:
426:
417:
416:
406:
397:(6): 1257–1264.
382:
373:
372:
360:
351:
350:
334:
324:
311:
310:
274:
193:Direct mechanism
103:pedunculopontine
53:circadian rhythm
1417:
1416:
1412:
1411:
1410:
1408:
1407:
1406:
1387:
1386:
1385:
1380:
1275:Procrastination
1228:Four-poster bed
1197:
1161:
1155:Polysomnography
1133:Sleep induction
1109:
1080:Sleep paralysis
1039:
991:
950:
947:
939:
881:
840:Mouth breathing
818:Sleep disorders
812:
749:
740:Quiescent sleep
720:
718:sleep disorders
711:
681:
680:
659:(12): 726–731.
649:
645:
600:
596:
561:
557:
526:
522:
499:
495:
463:
459:
427:
420:
383:
376:
361:
354:
347:
325:
314:
275:
266:
261:
244:
235:
195:
167:
159:cerebral cortex
131:locus coeruleus
119:locus coeruleus
91:cerebral cortex
87:basal forebrain
74:
65:
57:basal forebrain
40:
17:
12:
11:
5:
1415:
1405:
1404:
1399:
1382:
1381:
1379:
1378:
1373:
1368:
1359:
1354:
1349:
1344:
1339:
1334:
1329:
1324:
1319:
1314:
1309:
1304:
1302:Comfort object
1299:
1294:
1289:
1284:
1283:
1282:
1277:
1267:
1262:
1257:
1252:
1251:
1250:
1245:
1240:
1235:
1230:
1225:
1220:
1209:
1207:
1203:
1202:
1199:
1198:
1196:
1195:
1190:
1185:
1180:
1175:
1173:Sleep medicine
1169:
1167:
1163:
1162:
1160:
1159:
1158:
1157:
1147:
1146:
1145:
1140:
1130:
1125:
1119:
1117:
1111:
1110:
1108:
1107:
1102:
1097:
1092:
1087:
1082:
1077:
1072:
1063:
1058:
1053:
1047:
1045:
1041:
1040:
1038:
1037:
1032:
1027:
1022:
1017:
1012:
1007:
1001:
999:
993:
992:
990:
989:
984:
979:
974:
969:
964:
959:
953:
951:
944:
941:
940:
938:
937:
932:
927:
922:
917:
912:
907:
902:
897:
891:
889:
883:
882:
880:
879:
874:
873:
872:
867:
862:
857:
852:
842:
837:
831:
829:
820:
814:
813:
811:
810:
805:
800:
795:
790:
785:
780:
775:
770:
765:
759:
757:
751:
750:
748:
747:
742:
737:
731:
729:
722:
721:
710:
709:
702:
695:
687:
679:
678:
643:
614:(3): 977–990.
594:
575:(1): 269–292.
555:
536:(12): 726–31.
520:
493:
457:
438:(4): 1377–90.
418:
374:
352:
345:
312:
263:
262:
260:
257:
256:
255:
250:
243:
240:
234:
231:
194:
191:
166:
163:
73:
70:
64:
63:Neural circuit
61:
39:
36:
15:
9:
6:
4:
3:
2:
1414:
1403:
1400:
1398:
1395:
1394:
1392:
1377:
1374:
1372:
1369:
1367:
1363:
1360:
1358:
1355:
1353:
1350:
1348:
1345:
1343:
1340:
1338:
1335:
1333:
1330:
1328:
1325:
1323:
1320:
1318:
1315:
1313:
1310:
1308:
1305:
1303:
1300:
1298:
1295:
1293:
1290:
1288:
1285:
1281:
1278:
1276:
1273:
1272:
1271:
1268:
1266:
1263:
1261:
1258:
1256:
1253:
1249:
1246:
1244:
1241:
1239:
1236:
1234:
1231:
1229:
1226:
1224:
1221:
1219:
1216:
1215:
1214:
1211:
1210:
1208:
1204:
1194:
1191:
1189:
1186:
1184:
1181:
1179:
1176:
1174:
1171:
1170:
1168:
1164:
1156:
1153:
1152:
1151:
1148:
1144:
1141:
1139:
1136:
1135:
1134:
1131:
1129:
1128:Sleep hygiene
1126:
1124:
1121:
1120:
1118:
1116:
1112:
1106:
1103:
1101:
1098:
1096:
1093:
1091:
1088:
1086:
1085:Sleep inertia
1083:
1081:
1078:
1076:
1073:
1071:
1067:
1064:
1062:
1059:
1057:
1054:
1052:
1049:
1048:
1046:
1042:
1036:
1035:Sleep-talking
1033:
1031:
1030:Sleep driving
1028:
1026:
1023:
1021:
1018:
1016:
1013:
1011:
1008:
1006:
1003:
1002:
1000:
998:
994:
988:
985:
983:
980:
978:
975:
973:
970:
968:
965:
963:
960:
958:
955:
954:
952:
949:
942:
936:
933:
931:
928:
926:
923:
921:
918:
916:
913:
911:
908:
906:
903:
901:
898:
896:
893:
892:
890:
888:
884:
878:
875:
871:
868:
866:
863:
861:
858:
856:
853:
851:
848:
847:
846:
843:
841:
838:
836:
833:
832:
830:
828:
824:
821:
819:
815:
809:
806:
804:
803:Sleep spindle
801:
799:
796:
794:
791:
789:
786:
784:
781:
779:
776:
774:
771:
769:
766:
764:
761:
760:
758:
756:
752:
746:
743:
741:
738:
736:
733:
732:
730:
728:
723:
719:
715:
708:
703:
701:
696:
694:
689:
688:
685:
674:
670:
666:
662:
658:
654:
647:
639:
635:
630:
625:
621:
617:
613:
609:
605:
598:
590:
586:
582:
578:
574:
570:
566:
559:
551:
547:
543:
539:
535:
531:
524:
516:
512:
509:(3): 283–93.
508:
504:
497:
489:
485:
481:
477:
474:(3): 669–83.
473:
469:
461:
453:
449:
445:
441:
437:
433:
425:
423:
414:
410:
405:
400:
396:
392:
388:
381:
379:
370:
366:
359:
357:
348:
342:
338:
333:
332:
323:
321:
319:
317:
308:
304:
300:
296:
292:
288:
284:
280:
273:
271:
269:
264:
254:
251:
249:
246:
245:
239:
230:
226:
222:
220:
216:
212:
208:
204:
200:
190:
188:
184:
180:
176:
173:and anterior
172:
171:preoptic area
162:
160:
156:
155:acetylcholine
152:
148:
144:
140:
136:
135:noradrenaline
132:
128:
124:
120:
116:
112:
108:
104:
100:
99:acetylcholine
96:
92:
88:
84:
80:
69:
60:
58:
54:
49:
45:
35:
34:(REM sleep).
33:
29:
25:
21:
1248:Sleeping bag
1069:
1025:Sleepwalking
1010:Night terror
727:sleep cycles
656:
652:
646:
611:
607:
597:
572:
569:Neuroscience
568:
558:
533:
529:
523:
506:
502:
496:
471:
468:Neuroscience
467:
460:
435:
432:Neuroscience
431:
404:10.2741/1159
394:
390:
368:
364:
330:
282:
278:
236:
227:
223:
196:
175:hypothalamus
168:
139:raphe nuclei
123:raphe nuclei
83:hypothalamus
75:
66:
44:encephalitis
41:
19:
18:
1332:Second wind
1307:Dream diary
1183:Sleep study
1123:Sleep diary
1075:Hypnopompia
1070:Sleep onset
1061:Hypnic jerk
900:Hypersomnia
850:Catathrenia
845:Sleep apnea
755:Brain waves
725:Stages of
205:type A and
77:respective
20:Sleep onset
1391:Categories
1366:Sleep debt
1312:Microsleep
1292:Chronotype
1206:Daily life
1090:Somnolence
1066:Hypnagogia
997:Parasomnia
915:Narcolepsy
827:Anatomical
808:Theta wave
778:Gamma wave
773:Delta wave
763:Alpha wave
371:: 7:33–38.
259:References
253:Hypnagogia
211:NREM sleep
115:NREM sleep
28:narcolepsy
1376:Sleepover
1327:Power nap
1322:Nightwear
1188:Melatonin
1150:Somnology
1115:Treatment
948:disorders
887:Dyssomnia
793:PGO waves
788:Mu rhythm
783:K-complex
768:Beta wave
745:Slow-wave
111:REM sleep
1243:Mattress
1218:Bunk bed
1138:Hypnosis
930:Nocturia
905:Insomnia
673:11718878
638:11826126
589:19285545
550:11718878
452:16039802
413:12957869
307:16251950
242:See also
179:insomnia
177:lead to
133:produce
85:and the
72:Pathways
1270:Bedtime
1265:Bedroom
1260:Bedding
1255:Bed bug
1238:Hammock
1143:Lullaby
977:Jet lag
877:Snoring
835:Bruxism
629:6758527
515:5724423
488:2601839
287:Bibcode
233:Stage 1
207:galanin
165:Lesions
38:History
1337:Siesta
1223:Daybed
1051:Dreams
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279:Nature
147:orexin
1397:Sleep
1280:Story
1233:Futon
1166:Other
714:Sleep
716:and
669:PMID
634:PMID
585:PMID
546:PMID
511:PMID
484:PMID
448:PMID
409:PMID
341:ISBN
303:PMID
219:GABA
215:GABA
203:GABA
187:GABA
153:and
151:GABA
105:and
1317:Nap
1213:Bed
661:doi
624:PMC
616:doi
577:doi
573:161
538:doi
507:106
476:doi
440:doi
436:134
399:doi
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