Circadian and wakefulness-sleep modulation of cognition in humans

Frontiers in Molecular Neuroscience

Volumn , Issue APRIL, 2012, Pages

  Wright, Kenneth P ^a   Lowry, Christopher A ^a   leBourgeois, Monique K ^a  

^a UNIVERSITY OF COLORADO   (United States)

Author keywords

Circadian phase; Circadian sleep disorders; Forced desynchrony; Performance; Sleep homeostasis

Indexed keywords

AROUSAL; BIOLOGICAL RHYTHM; CIRCADIAN MISALIGNMENT; CIRCADIAN RHYTHM; CIRCADIAN RHYTHM SLEEP DISORDER; COGNITION; COGNITIVE DEFECT; EMOTIONAL DISORDER; ENVIRONMENTAL STRESS; HUMAN; JET LAG; LEARNING DISORDER; REM SLEEP; REVIEW; SLEEP DEPRIVATION; SLEEP PARAMETERS; SLEEP PATTERN; SLEEP PROMOTING SYSTEM; SLEEP TIME; SLEEP WAKING CYCLE; SUPRACHIASMATIC NUCLEUS;

EID: 84864102922     PISSN: 16625099     EISSN: None     Source Type: Journal    
DOI: None     Document Type: Review

References (166)

1. Abe, M., Herzog, E. D., Yamazaki, S., Straume, M., Tei, H., Sakaki, Y., Menaker, M., and Block, G. D. (2002). Circadian rhythms in isolated brain regions. J. Neurosci. 22, 350-356.
2. Abrams, J. K., Johnson, P. L., Hollis, J. H., and Lowry, C. A. (2004). Anatomic and functional topography of the dorsal raphe nucleus. Ann. N.Y. Acad. Sci. 1018, 46-57.
3. Åkerstedt, T. (1998). Shift work and disturbed sleep/wakefulness. Sleep Med. Rev. 2, 117-128.
4. Åkerstedt, T., Fredlund, P., Gillberg, M., and Jansson, B. (2002). Work load and work hours in relation to disturbed sleep and fatigue in a large representative sample. J. Psychosom. Res. 53, 585-588.
5. Alam, M. N., Szymusiak, R., Gong, H., King, J., and McGinty, D. (1999). Adenosinergic modulation of rat basal forebrain neurons during sleep and waking: neuronal recording with microdialysis. J. Physiol. 521, 679-690.
6. Aston-Jones, G., and Bloom, F. E. (1981). Activity of norepinephrine-containing locus coeruleus neurons in behaving rats anticipates fluctuations in the sleep-waking cycle. J. Neurosci. 1, 876-886.
7. Aston-Jones, G., Chen, S., Zhu, Y., and Oshinsky, M. L. (2001). A neural circuit for circadian regulation of arousal. Nat. Neurosci. 4, 732-738.
8. Aston-Jones, G., Shipley, M. T., Chouvet, G., Ennis, M., van Bockstaele, E., Pieribone, V., Shiekhattar, R., Akaoka, H., Drolet, G., Astier, B., Charléty, P., Valentino, R. J., and Williams, J. T. (1991). Afferent regulation of locus coeruleus neurons: anatomy, physiology and pharmacology. Prog. Brain Res. 88, 47-75.
9. Balkin, T. J., Braun, A. R., Wesensten, N. J., Jeffries, K., Varga, M., Baldwin, P., Belenky, G., and Herscovitch, P. (2002). The process of awakening: a PET study of regional brain activity patterns mediating the reestablishment of alertness and consciousness. Brain 125, 2308-2319.
10. Barger, L. K., Cade, B. E., Ayas, N. T., Cronin, J. W., Rosner, B., Speizer, F. E., Czeisler, C. A., and Harvard Work Hours, Health, and Safety Group. (2005). Extended work shifts and the risk of motor vehicle crashes among interns. N. Engl. J. Med. 352, 125-134.
11. Bayer, L., Eggermann, E., Serafin, M., Grivel, J., Machard, D., Muhlethaler, M., and Jones, B. E. (2005). Opposite effects of noradrenaline and acetylcholine upon hypocretin/orexin versus melanin concentrating hormone neurons in rat hypothalamic slices. Neuroscience 130, 807-811.
12. Boivin, D. B., Czeisler, C. A., Dijk, D. J., Duffy, J. F., Folkard, S., Minors, D. S., Totterdell, P., and Waterhouse, J. M. (1997). Complex interaction of the sleep-wake cycle and circadian phase modulates mood in healthy subjects. Arch. Gen. Psychiatry 54, 145-152.
13. Brabant, C., Alleva, L., Quertemont, E., and Tirelli, E. (2010). Involvement of the brain histaminergic system in addiction and addiction-related behaviors: a comprehensive review with emphasis on the potential therapeutic use of histaminergic compounds in drug dependence. Prog. Neurobiol. 92, 421-441.
14. Brown, R. E., Sergeeva, O. A., Eriksson, K. S., and Haas, H. L. (2002). Convergent excitation of dorsal raphe serotonin neurons by multiple arousal systems (orexin/hypocretin, histamine and noradrenaline). J. Neurosci. 22, 8850-8859.
15. Brown, R. E., Stevens, D. R., and Haas, H. L. (2001). The physiology of brain histamine. Prog. Neurobiol. 63, 637-672.
16. Buhr, E. D., Yoo, S. H., and Takahashi, J. S. (2010). Temperature as a universal resetting cue for mammalian circadian oscillators. Science 330, 379-385.
17. Carskadon, M. A., Harvey, K., Duke, P., Anders, T. F., Litt, I. F., and Dement, W. C. (1980). Pubertal changes in daytime sleepiness. Sleep 2, 453-460.
18. Carskadon, M. A., Wolfson, A. R., Acebo, C., Tzischinsky, O., and Seifer, R. (1998). Adolescent sleep patterns, circadian timing, and sleepiness at a transition to early school days. Sleep 21, 871-881.
19. Cermakian, N., Lamont, E. W., Boudreau, P., and Boivin, D. B. (2011). Circadian clock gene expression in brain regions of alzheimer's disease patients and control subjects. J. Biol. Rhythms26, 160-170.
20. Chamberlin, N. L., Arrigoni, E., Chou, T. C., Scammell, T. E., Greene, R. W., and Saper, C. B. (2003). Effects of adenosine on gabaergic synaptic inputs to identified ventrolateral preoptic neurons. Neuroscience 119, 913-918.
21. Chou, T. C., Bjorkum, A. A., Gaus, S. E., Lu, J., Scammell, T. E., and Saper, C. B. (2002). Afferents to the ventrolateral preoptic nucleus. J. Neurosci. 22, 977-990.
22. Chou, T. C., Scammell, T. E., Gooley, J. J., Gaus, S. E., Saper, C. B., and Lu, J. (2003). Critical role of dorsomedial hypothalamic nucleus in a wide range of behavioral circadian rhythms. J. Neurosci. 23, 10691-10702.
23. Cohen, D. A., Wang, W., Wyatt, J. K., Kronauer, R. E., Dijk, D. J., Czeisler, C. A., and Klerman, E. B. (2010). Uncovering residual effects of chronic sleep loss on human performance. Sci. Transl. Med. 2, 14ra3.
24. Crosby, B., Gryczkowski, M., LeBourgeois, M. K., Olmi, D., Rabian, B., and Harsh, J. R. (2006). Mid-sleep time and school readiness in black and white preschool children. Sleep 29, A78.
25. Crosby, B., LeBourgeois, M. K., and Harsh, J. (2005). Racial differences in reported napping and nocturnal sleep in 2-to 8-year-old children. Pediatrics 115, 225-232.
26. Czeisler, C. A., Dijk, D.-J., and Duffy, J. F. (1994). "Entrained phase of the circadian pacemaker serves to stabilize alertness and performance throughout the habitual waking day. " in Sleep Onset: Normal and Abnormal Processes, eds R. D. Ogilvie and J. R. Harsh (Washington, DC: American Psychological Association), 89-110.
27. Czeisler, C. A., Walsh, J. K., Roth, T., Hughes, R. J., Wright, K. P., Kingsbury, L., Arora, S., Schwartz, J. R., Niebler, G. E., Dinges, D. F., and U.S. Modafinil in Shift Work Sleep Disorder Study Group. (2005). Modafinil for excessive sleepiness associated with shift-work sleep disorder. N. Engl. J. Med. 353, 476-486.
28. Dagan, Y., Stein, D., Steinbock, M., Yovel, I., and Hallis, D. (1998). Frequency of delayed sleep phase syndrome among hospitalized adolescent psychiatric patients. J. Psychosom. Res. 45, 15-20.
29. Dai, J., Swaab, D. F., and Buijs, R. M. (1997). Distribution of vasopressin and vasoactive intestinal polypeptide (VIP) fibers in the human hypothalamus with special emphasis on suprachiasmatic nucleus efferent projections. J. Comp. Neurol. 383, 397-414.
30. Date, Y., Ueta, Y., Yamashita, H., Yamaguchi, H., Matsukura, S., Kangawa, K., Sakurai, T., Yanagisawa, M., and Nakazato, M. (1999). Orexins, orexigenic hypothalamic peptides, interact with autonomic, neuroendocrine and neuroregulatory systems. Proc. Natl. Acad. Sci. U.S.A. 96, 748-753.
31. Datta, S., and Siwek, D. F. (2002). Single cell activity patterns of pedunculopontine tegmentum neurons across the sleep-wake cycle in the freely moving rats. J. Neurosci. Res. 70, 611-621.
32. de Lecea, L., Kilduff, T. S., Peyron, C., Gao, X., Foye, P. E., Danielson, P. E., Fukuhara, C., Battenberg, E. L., Gautvik, V. T., Bartlett, F. S. 2nd, Frankel, W. N., van den Pol, A. N., Bloom, F. E., Gautvik, K. M., and Sutcliffe, J. G. (1998). The hypocretins: hypothalamus-specific peptides with neuroexcitatory activity. Proc. Natl. Acad. Sci. U.S.A. 95, 322-327.
33. Deurveilher, S., and Semba, K. (2003). Indirect projections from the suprachiasmatic nucleus to the median preoptic nucleus in rat. Brain Res. 987, 100-106.
34. Deurveilher, S., and Semba, K. (2005). Indirect projections from the suprachiasmatic nucleus to major arousal-promoting cell groups in rat: implications for the circadian control of behavioural state. Neuroscience 130, 165-183.
35. Dijk, D. J., and Czeisler, C. A. (1995). Contribution of the circadian pacemaker and the sleep homeostat to sleep propensity, sleep structure, electroencephalographic slow waves, and sleep spindle activity in humans. J. Neurosci. 15, 3526-3538.
36. Dinges, D. F. (1990). "Are you awake? Cognitive performance and reverie during the hypnopom-pic state. " in Sleep And Cognition, eds R. Bootzin, J. Kihlstrom, and D. Schacter (Washington, DC: American Psychological Association), 159-175.
37. Dinges, D. F. (1995). An overview of sleepiness and accidents. J. Sleep Res. 4, 4-14.
38. Drake, C. L., Roehrs, T., Richardson, G., Walsh, J. K., and Roth, T. (2004). Shift work sleep disorder: prevalence and consequences beyond that of symptomatic day workers. Sleep 27, 1453-1462.
39. Drake, C. L., and Wright, K. P. Jr. (2011). "Shift work, shift work disorder, and jet lag, " in Principles and Practice of Sleep Medicine, eds M. H. Kryger, T. Roth, and W. C. Dement 5 edn., (St. Louis: Saunders), 784-798.
40. Jones, J. F., Cain, S. W., Chang, A. M., Phillips, A. J., Münch, M. Y., Gronfier, C., Wyatt, J. K., Dijk, D. J., Wright, K. P. Jr., and Czeisler, C. A. (2011). Quantification of behavior sackler colloquium: sex difference in the near-24-hour intrinsic period of the human circadian timing system. Proc. Natl. Acad. Sci. U.S.A. 108, 15602-15608.
41. Duffy, J. F., Dijk, D. J., Hall, E. F., and Czeisler, C. A. (1999). Relationship of endogenous circadian melatonin and temperature rhythms to selfreported preference for morning or evening activity in young and older people. J. Investig. Med. 47, 141-150.
42. Duffy, J. F., Rimmer, D. W., and Czeisler, C. A. (2001). Association of intrinsic circadian period with morningness-eveningness, usual wake time, and circadian phase. Behav. Neurosci. 115, 895-899.
43. Dumont, M., Benhaberou-Brun, D., and Paquet, J. (2001). Profile of 24-h light exposure and circadian phase of melatonin secretion in night workers. J. Biol. Rhythms 16, 502-511.
44. Ebisawa, T., Uchiyama, M., Kajimura, N., Mishima, K., Kamei, Y., Katoh, M., Watanabe, T., Sekimoto, M., Shibui, K., Kim, K., Kudo, Y., Ozeki, Y., Sugishita, M., Toyoshima, R., Inoue, Y., Yamada, N., Nagase, T., Ozaki, N., Ohara, O., Ishida, N., Okawa, M., Takahashi, K., and Yamauchi, T. (2001). Association of structural polymorphisms in the human period3 gene with delayed sleep phase syndrome. EMBO Rep. 2, 342-346.
45. Elmenhorst, D., Meyer, P. T., Winz, O. H., Matusch, A., Ermert, J., Coenen, H. H., Basheer, R., Haas, H. L., Zilles, K., and Bauer, A. (2007). Sleep deprivation increases A1 adenosine receptor binding in the human brain: a positron emission tomography study. J. Neurosci. 27, 2410-2415.
46. Frey, D. J., Fleshner, M., and Wright, K. P. Jr. (2007). The effects of 40 hours of total sleep deprivation on inflammatory markers in healthy young adults. Brain Behav. Immun. 21, 1050-1057.
47. Fuller, P. M., Saper, C. B., and Lu, J. (2007). The pontine REM switch: past and present. J. Physiol. 584, 735-741.
48. Garlo, K. G., Crossin, R. A., Carskadon, M. A., and LeBourgeois, M. K. (2008). Assessment of salivary dim light melatonin onset (DLMO) and reported sleep in preschool children. Sleep 31, A48.
49. Goel, N., Banks, S., Mignot, E., and Dinges, D. F. (2009). PER3 polymorphism predicts cumulative sleep homeostatic but not neurobehav-ioral changes to chronic partial sleep deprivation. PLoS One 4:e5874. doi: 10.1371/journal.pone.0005874
50. Grady, S., Aeschbach, D., Wright, K. P. Jr., and Czeisler, C. A. (2010). Effect of modafinil on impairments in neurobehavioral performance and learning associated with extended wakefulness and circadian misalignment. Neuropsychopharmacolgy 35, 1910-1920.
51. Greene, C. M., Bellgrove, M. A., Gill, M., and Robertson, I. H. (2009). Noradrenergic genotype predicts lapses in sustained attention. Neuropsychologia 47, 591-594.
52. Gronfier, C., Wright, K. P. Jr., Kronauer, R. E., and Czeisler, C. A. (2007). Entrainment of the human circadian pacemaker to longer-than-24-h days. Proc. Natl. Acad. Sci. U.S.A. 104, 9081-9086.
53. Gooley, J. J., Lu, J., Fischer, D., and Saper, C. B. (2003). A broad role for melanopsin in nonvisual photoreception. J. Neurosci. 23, 7093-106.
54. Guilding, C., Hughes, A. T., Brown, T. M., Namvar, S., and Piggins, H. D. (2009). A riot of rhythms: neuronal and glial circadian oscillators in the mediobasal hypothalamus. Mol. Brain 2, 28.
55. Guilding, C., and Piggins, H. D. (2007). Challenging the omnipotence of the suprachiasmatic timekeeper: are circadian oscillators present throughout the mammalian brain? Eur. J. Neurosci. 25, 3195-3216.
56. Hagan, J. J., Leslie, R. A., Patel, S., Evans, M. L., Wattam, T. A., Holmes, S., Benham, C. D., Taylor, S. G., Routledge, C., Hemmati, P., Munton, R. P., Ashmeade, T. E., Shah, A. S., Hatcher, J. P., Hatcher, P. D., Jones, D. N., Smith, M. I., Piper, D. C., Hunter, A. J., Porter, R. A., and Upton, N. (1999). Orexin A activates locus coeruleus cell firing and increases arousal in the rat. Proc. Natl. Acad. Sci. U.S.A. 96, 10911-10916.
57. Hale, M. W., and Lowry, C. A. (2011). Functional topography of midbrain and pontine serotonergic systems: implications for synaptic regulation of serotonergic circuits. Psychopharmacol 213, 243-264.
58. Hannibal, J., and Fahrenkrug, J. (2004). Target areas innervated by PACAP-immunoreactive retinal ganglion cells. Cell Tissue Res. 316, 99-113.
59. Hassani, O. K., Henny, P., Lee, M. G., and Jones, B. E. (2010). GABAergic neurons intermingled with orexin and MCH neurons in the lateral hypothalamus discharge maximally during sleep. Eur. J. Neurosci. 32, 448-457.
60. Hayakawa, T., Uchiyama, M., Kamei, Y., Shibui, K., Tagaya, H., Asada, T., Okawa, M., Urata, J., and Takahashi, K. (2005). Clinical analyses of sighted patients with non-24-hour sleep-wake syndrome: a study of 57 consecutively diagnosed cases. Sleep 28, 945-952.
61. Henny, P., and Jones, B. E. (2006). Innervation of orexin/hypocretin neurons by GABAergic, glutamater-gic or cholinergic basal forebrain terminals evidenced by immunos-taining for presynaptic vesicular transporter and postsynaptic scaffolding proteins. J. Comp. Neurol. 499, 645-661.
62. Hohjoh, H., Takasu, M., Shishikura, K., Takahashi, Y., Honda, Y., and Tokunaga, K. (2003). Significant association of the arylalkylamine N-acetyltransferase (AA-NAT) gene with delayed sleep phase syndrome. Neurogenetics 4, 151-153.
63. Horne, J. A., and Reyner, L. A. (1995). Sleep related vehicle accidents. BMJ 310, 565-567.
64. Horne, J. A., Reyner, L. A., and Barrett, P. R. (2003). Driving impairment due to sleepiness is exacerbated by low alcohol intake. Occup. Environ. Med. 60, 689-692.
65. Hsieh, K. C., Gvilia, I., Kumar, S., Uschakov, A., McGinty, D., Alam, M. N., and Szymusiak, R. (2011). c-Fos expression in neurons projecting from the preoptic and lateral hypothalamic areas to the ventrolateral periaqueductal gray in relation to sleep states. Neuroscience 188, 55-67.
66. Hull, J. T., Wright, K. P., and Czeisler, C. A. (2003). The influence of subjective alertness and motivation on human performance independent of circadian and homeostatic regulation. J. Biol. Rhythms 18, 329-338.
67. Hughes, A. T., Guilding, C., and Piggins, H. D. (2011). Neuropeptide signaling differentially affects phase maintenance and rhythm generation in SCN and extra-SCN circadian oscillators. PLoS One 6:e18926. doi: 10.1371/journal.pone.0018926
68. Iglowstein, I., Jenni, O. G., Molinari, L., and Largo, R. H. (2003). Sleep duration from infancy to adolescence: reference values and generational trends. Pediatrics 111, 302-307.
69. Jenni, O. G., Achermann, P., and Carskadon, M. A. (2005). Homeostatic sleep regulation in adolescents. Sleep28, 1446-1454.
70. Jenni, O. G., and LeBourgeois, M. K. (2006). Understanding sleep-wake behavior and sleep disorders in children: the value of a model. Curr. Opin. Psychiatry19, 282-287.
71. Jilg, A., Lesny, S., Peruzki, N., Schwegler, H., Selbach, O., Dehghani, F., and Stehle, J. H. (2010). Temporal dynamics of mouse hippocampal clock gene expression support memory processing. Hippocampus 20, 377-388.
72. Johnson, P. L., Truitt, W., Fitz, S. D., Minick, P. E., Dietrich, A., Sanghani, S., Träskman-Bendz, L., Goddard, A. W., Brundin, L., and Shekhar, A. (2010). A key role for orexin in panic anxiety. Nat. Med. 16, 111-149.
73. Jones, B. E. (2005). From waking to sleeping: neuronal and chemical substrates. Trends Pharmacol. Sci. 26, 578-586.
74. Jones, B. E. (2008). Modulation of cortical activation and behavioral arousal by cholinergic and orexin-ergic systems. Ann. N.Y. Acad. Sci. 1129, 26-34.
75. Kalsbeek, A., Fliers, E., Hofman, M. A., Swaab, D. F., and Buijs, R. M. (2010). Vasopressin and the output of the hypothalamic biological clock. J. Neuroendocrinol. 22, 362-372.
76. Karatsoreos, I. N., Bhagat, S., Bloss, E. B., Morrison, J. H., and McEwen, B. S. (2011). Disruption of circadian clocks has ramifications for metabolism, brain, and behavior. Proc. Natl. Acad. Sci. U.S.A. 108, 1657-1662.
77. Katz, G. (2011). Jet lag and psychotic disorders. Curr. Psychiatry Rep. 13, 187-192.
78. Katz, G., Durst, R., Zislin, Y., Barel, Y., and Knobler, H. Y. (2001). Psychiatric aspects of jet lag: review and hypothesis. Med. Hypotheses56, 20-23.
79. Kerkhof, G. A., and van Dongen, H. P. (1996). Morning-type and evening-type individuals differ in the phase position of their endogenous circadian oscillator. Neurosci. Lett. 218, 153-156.
80. Kohler, C. A., da Silva, W. C., Benetti, F., and Bonini, J. S. (2011). Histaminergic mechanisms for modulation of memory systems. Neural. Plast. 2011, 328602.
81. Krauchi, K., Cajochen, C., and Wirz-Justice, A. (1997). A relationship between heat loss and sleepiness: effects of postural change and melatonin administration. J. Appl. Physiol. 83, 134-139.
82. Krueger, J. M., Clinton, J. M., Winters, B. D., Zielinski, M. R., Taishi, P., Jewett, K. A., and Davis, C. J. (2011). Involvement of cytokines in slow wave sleep. Prog. Brain Res. 193, 39-47.
83. Landrigan, C. P., Rothschild, J. M., Cronin, J. W., Kaushal, R., Burdick, E., Katz, J. T., Lilly, C. M., Stone, P. H., Lockley, S. W., Bates, D. W., and Czeisler, C. A. (2004). Effect of reducing interns' work hours on serious medical errors in intensive care units. N. Engl. J. Med. 351, 1838-1848.
84. Leak, R. K., and Moore, R. Y. (2001). Topographic organization of suprachiasmatic nucleus projection neurons. J. Comp. Neurol. 433, 312-334.
85. Lee, J. H., Wang, W., Silva, E. J., Chang, A. M., Scheuermaier, K. D., Cain, S. W., and Duffy, J. F. (2009). Neurobehavioral performance in young adults living on a 28-h day for 6 weeks. Sleep 32, 905-913.
86. Li, S. X., Liu, L. J., Jiang, W. G., and Lu, L. (2009). Morphinewithdrawal produces circadian rhythm alterations of clock genes in mesolimbic brain areas and peripheral blood mononuclear cells in rats. J. Neurochem. 109, 1668-2679.
87. Liu, R. J., van den Pol, A. N., and Aghajanian, G. K. (2002). Hypocretins (orexins) regulate serotonin neurons in the dorsal raphe nucleus by excitatory direct and inhibitory indirect actions. J. Neurosci. 22, 9453-9464.
88. Lockley, S. W., Cronin, J. W., Evans, E. E., Cade, B. E., Lee, C. J., Landrigan, C. P., Rothschild, J. M., Katz, J. T., Lilly, C. M., Stone, P. H., Aeschbach, D., and Czeisler, C. A. Harvard Work Hours, Health, and Safety Group. (2004). Effect of reducing interns' weeklywork hours on sleep and attentional failures. N. Engl. J. Med. 351, 1829-1837.
89. Lowry, C. A., Evans, A. K., Gasser, P. J., Hale, M. W., Staub, D. R., and Shekhar, A. (2008). "Topographical organization and chemoarchitec-ture of the dorsal raphe nucleus and the median raphe nucleus, " in: Serotonin and Sleep: Molecular, Functional and Clinical Aspects, eds J. M. Monti, B. L. Pandi-Perumal, B. L. Jacobs, and D. L. Nutt (Basel: Birkhauser), 25-68. F. Koob, M. Le Moal, and R. F. Thompson (Oxford: Academic Press), 295-303).
90. Lu, B. S., and Zee, P. C. (2006). Circadian rhythm sleep disorders. Chest 130, 1915-1923.
91. Lu, J., Greco, M. A., Shiromani, P., and Saper, C. B. (2000). Effect of lesions of the ventrolateral preoptic nucleus on NREM and REM sleep. J. Neurosci. 20, 3830-3842.
92. Luo, A. H., and Aston-Jones, G. (2009). Circuit projection from suprachias-matic nucleus to ventral tegmental area: a novel circadian output pathway. Eur. J. Neurosci. 29, 748-760.
93. Lyznicki, J. M., Doege, T. C., Davis, R. M., and Williams, M. A. (1998). Sleepiness, driving, and motor vehicle crashes. JAMA 279, 1908-1913.
94. Markwald, R. R., Lee-Chiong, T. L., Burke, T. M., Snider, J. A., and Wright, K. P. Jr. (2010). Effects of the melatonin MT-1/MT-2 agonist ramelteon on daytime body temperature and sleep. Sleep 33, 825-831.
95. Masubuchi, S., Honma, S., Abe, H., Ishizaki, K., Namihira, M., Ikeda, M., and Honma, K. (2000). Clock genes outside the suprachiasmatic nucleus involved in manifestation of locomotor activity rhythm in rats. Eur. J. Neurosci. 12, 4206-4214.
96. Matthews, R. W., Ferguson, S.A., Zhou, X., Kosmadopoulos, A., Kennaway, D. J., and Roach, G. D. (2012). Simulated driving under the influence of extended wake, time of day and sleep restriction. Accid. Anal. Prev. 45, 55-61.
97. McCarley, R. W. (2007). Neurobiology of REM and NREM sleep. Sleep Med. 8, 302-330.
98. McCormick, D. A., and Bal, T. (1997). Sleep and arousal: thalamocortical mechanisms. Annu. Rev. Neurosci. 20, 185-215.
99. Mongrain, V., Lavoie, S., Selmaoui, B., Paquet, J., and Dumont, M. (2004). Phase relationships between sleep-wake cycle and underlying circadian rhythms in morningness-eveningness. J. Biol. Rhythms 19, 248-257.
100. Monti, J. M., and Monti, D. (2007). The involvement of dopamine in the modulation ofsleep and waking. Sleep Med. Rev. 11, 113-133.
101. Murray, G., Nicholas, C. L., Kleiman, J., Dwyer, R., Carrington, M. J., Allen, N. B., and Trinder, J. (2009). Nature's clocks and human mood: the circadian system modulates reward motivation. Emotion 9, 705-716.
102. Namihira, M., Honma, S., Abe, H., Tanahashi, Y., Ikeda, M., and Honma, K. (1999). Daily variation and light responsiveness of mammalian clock gene, Clock and BMAL1, transcripts in the pineal body and different areas of brain in rats. Neurosci. Lett. 267, 69-72.
103. Nitz, D., and Siegel, J. (1997). GABA release in the dorsal raphe nucleus: role in the control of REM sleep. Am. J. Physiol. 273, R451-R455.
104. Okawa, M., and Uchiyama, M. (2007). Circadian rhythm sleep disorders: characteristics and entrainment pathology in delayed sleep phase and non-24 sleep-wake syndrome. Sleep Med. Rev. 11, 485-496.
105. Owens, J. A., Belon, K., and Moss, P. (2010). Impact of delaying school start time on adolescent sleep, mood, and behavior. Arch. Pediatr. Adolesc. Med. 164, 608-614.
106. Pack, A. I., Pack, A. M., Rodgman, E., Cucchiara, A., Dinges, D. F., and Schwab, C. W. (1995). Characteristics of crashes attributed to the driver having fallen asleep. Accid. Anal. Prev. 27, 769-775.
107. Passani, M. B., Blandina, P., and Torrealba, F. (2011). The histamine H3 receptor and eating behavior. J. Pharmacol. Exp. Ther. 336, 24-29.
108. Peyron, C., Tighe, D. K., van den Pol, A. N., de Lecea, L., Heller, H. C., Sutcliffe, J. G., and Kilduff, T. S. (1998). Neurons containing hypocretin (orexin) project to multiple neuronal systems. J. Neurosci. 18, 9996-10015.
109. Porkka-Heiskanen, T., Strecker, R. E., Thakkar, M., Bjorkum, A. A., Greene, R. W., and McCarley, R. W. (1997). Adenosine: a mediator of the sleep-inducing effects of prolonged wakefulness. Science 276, 1265-1268.
110. Rai, S., Kumar, S., Alam, M. A., Szymusiak, R., McGinty, D., and Alam, M. N. (2010). A(1) receptor mediated adenosinergic regulation of perifornical-lateral hypothalamic area neurons in freely behaving rats. Neuroscience 167, 40-48.
111. Rainnie, D. G., Grunze, H. C., McCarley, R. W., and Greene, R. W. (1994). Adenosine inhibition of mesopontine cholinergic neurons: implications for EEG arousal. Science 263, 689-692.
112. Rajaratnam, S. M., Polymeropoulos, M. H., Fisher, D. M., Roth, T., Scott, C., Birznieks, G., and Klerman, E. B. (2009a). Melatonin agonist tasimelteon (VEC-162) for transient insomnia after sleep-time shift: two randomised controlled multicentre trials. Lancet 373, 482-491.
113. Rajaratnam, S. M. W., Choen, D. A., and Rogers, N. L. (2009b). Melatonin and melatonin analogues. Sleep Med. Clin. 4, 179-193.
114. Reid, K. J., and Zee, P. C. (2009). Circadian rhythm disorders. Semin. Neurol. 29, 393-405.
115. Rosenwasser, A. M. (2010). Circadian clock genes: non-circadian roles in sleep, addiction, and psychiatric disorders? Neurosci. Biobehav. Rev. 34, 1249-1255.
116. Sack, R. L., Auckley, D., Auger, R. R., Carskadon, M. A., Wright, K. P. Jr., Vitiello, M. V., Zhdanova, I. V., and American Academy of Sleep Medicine. (2007a). Circadian rhythm sleep disorders: part i, basic principles, shift work and jet lag disorders: an american academy of sleep medicine review. Sleep 30, 1456-1479.
117. Sack, R. L., Auckley, D., Auger, R. R., Carskadon, M. A., Wright, K. P. Jr., Vitiello, M. V., Zhdanova, I. V., and American Academy of Sleep Medicine. (2007b). Circadian rhythm sleep disorders: part ii, advanced sleep phase disorder, delayed sleep phase disorder, free-running disorder, and irregular sleep-wake rhythm: an american academy of sleep medicine review. Sleep 30, 1480-1497.
118. Sack, R. L., Blood, M. L., and Lewy, A. J. (1992). Melatonin rhythms in night shiftworkers. Sleep 15, 434-441.
119. Sakai, K., and Crochet, S. (2001). Differentiation of presumed serotonergic dorsal raphe neurons in relation to behavior and wake-sleep states. Neuroscience 104, 1141-1155.
120. Saper, C. B., Fuller, P. M., Pedersen, N. P., Lu, J., andScammell, T. E. (2010). Sleep state switching. Neuron 68, 1023-1042.
121. Saper, C. B., Lu, J., Chou, T. C., and Gooley, J. (2005). The hypothalamic integrator for circadian rhythms. Trends Neurosci. 28, 152-157.
122. Saper, C. B., Scammell, T. E., and Lu, J. (2005). Hypothalamic regulation of sleep and circadian rhythms. Nature 437, 1257-1263.
123. Scheer, F. A., Shea, T. J., Hilton, M. F., and Shea, S. A. (2008). An endogenous circadian rhythm in sleep inertia results in greatest cognitive impairment upon awakening during the biological night. J. Biol. Rhythms 23, 353-361.
124. Schepens, B., Stapley, P., and Drew, T. (2008). Neurons in the pon-tomedullary reticular formation signal posture and movement both as an integrated behavior and independently. J. Neurophysiol. 100, 2235-2253.
125. Shen, H., and Semba, K. A. (1994). Direct retinal projection to the dorsal raphe nucleus in the rat. Brain Res. 635, 159-68.
126. Shouse, M. N., Staba, R. J., Saquib, S. F., and Farber, P. R. (2000). Monoamines and sleep: microdialysis findings in pons and amygdala. Brain Res. 860, 181-189.
127. Silva, E. J., and Duffy, J. F. (2008). Sleep inertia varies with circadian phase and sleep stage in older adults. Behav. Neurosci. 122, 928-935.
128. Silva, E. J., Wang, W., Ronda, J. M., Wyatt, J. K., and Duffy, J. F. (2010). Circadian and wake-dependent influences on subjective sleepiness, cognitive throughput, and reaction time performance in older and young adults. Sleep 33, 481-490.
129. Soffin, E. M., Gill, C. H., Brough, S. J., Jerman, J. C., and Davies, C. H. (2004). Pharmacological characterisation of the orexin receptor subtype mediating postsynaptic excitation in the rat dorsal raphe nucleus. Neuropharmacolgy 46, 1168-1176.
130. Steriade, M. (1992). Basic mechanisms of sleep generation. Neurol 42, 9-17.
131. Stiles, J., and Jernigan, T. L. (2010). The basics of brain development. Neuropsychol. Rev. 20, 327-348.
132. Strecker, R. E., Morairty, S., Thakkar, M. M., Porkka-Heiskanen, T., Basheer, R., Dauphin, L. J., Rainnie, D. G., Portas, C. M., Greene, R. W., and McCarley, R. W. (2000). Adenosinergic modulation of basal forebrain and preoptic/anterior hypothalamic neuronal activity in the control of behavioral state. Behav. Brain Res. 115, 183-204.
133. Szymusiak, R., Gvilia, I., and McGinty, D. (2007). Hypothalamic control of sleep. Sleep Med. 8, 291-301.
134. Szymusiak, R., and McGinty, D. (2008). Hypothalamic regulation of sleep and arousal. Ann. N.Y. Acad. Sci. 1129, 275-286.
135. Takahashi, K., Wang, Q. P., Guan, J. L., Kayama, Y., Shioda, S., and Koyama, Y. (2005). State-dependent effects of orexins on the serotonergic dorsal raphe neurons in the rat. Regul. Pept. 126, 43-47.
136. Takano, A., Uchiyama, M., Kajimura, N., Mishima, K., Inoue, Y., Kamei, Y., Kitajima, T., Shibui, K., Katoh, M., Watanabe, T., Hashimotodani, Y., Nakajima, T., Ozeki, Y., Hori, T., Yamada, N., Toyoshima, R., Ozaki, N., Okawa, M., Nagai, K., Takahashi, K., Isojima, Y., Yamauchi, T., and Ebisawa, T. (2004). A missense variation in human casein kinase I epsilon gene that induces functional alteration and shows an inverse association with circadian rhythm sleep disorders. Neuropsychopharmacology 29, 1901-1909.
137. Teclemariam-Mesbah, R., Ter Horst, G. J., Postema, F., Wortel, J., and Buijs, R. M. (1999). Anatomical demonstration of the suprachias-matic nucleus-pineal pathway. J. Comp. Neurol. 406, 171-182.
138. Toh, K. L., Jones, C. R., He, Y., Eide, E. J., Hinz, W. A., Virshup, D. M., Ptácek, L. J., and Fu, Y. H. (2001). An hPer2 phosphorylation site mutation in familial advanced sleep phase syndrome. Science 291, 1040-1043.
139. Tully, K., and Bolshakov, V. Y. (2010). Emotional enhancement of memory: how norepinephrine enables synaptic plasticity. Mol. Brain. 3, 15.
140. Uchiyama, M., and Lockley, S. W. (2009). Non-24-hour sleep-wake syndrome in sighted and blind patients. SleepMed. Clin. 4, 195-211.
141. Uschakov, A., Gong, H., McGinty, D., and Szymusiak, R. (2007). Efferent projections from the median preoptic nucleus to sleep-and arousal-regulatory nuclei in the rat brain. Neuroscience 150, 104-120.
142. Valdez, P., Ramirez, C., Garcia, A., Talamantes, J., and Cortez, J. (2010). Circadian and homeostatic variation in sustained attention. Chronobiol. Int. 27, 393-416.
143. van der Heijden, K. B., de Sonneville, L. M. J., and Althaus, M. (2010). Time-of-day effects on cognition in preadolescents: a trails study. Chronobiol. Int. 27, 1870-1894.
144. van Someren, E. J. W. (2000). More than a marker: interaction between the circadian regulation of temperature and sleep, age-related changes, and treatment possibilities. Chronobiol. Int. 17, 313-354.
145. Vgontzas, A. N., Zoumakis, E., Bixler, E. O., Lin, H. M., Follett, H., Kales, A., and Chrousos, G. P. (2004). Adverse effects of modest sleep restriction on sleepiness, performance, and inflammatory cytokines. J. Clin. Endocrinol. Metab. 89, 2119-2126.
146. Viola, A. U., Archer, S. N., James, L. M., Groeger, J. A., Lo, J. C., Skene, D. J., von Schantz, M., and Dijk, D. J. (2007). PER3 polymorphism predicts sleep structure and waking performance. Curr. Biol. 17, 613-618.
147. Vorona, R. D., Szklo-Coxe, M., Wu, A., Dubik, M., Zhao, Y. Q., and Ware, J. C. (2011). Dissimilar teen crash rates in two neighboring southeastern virginia cities with different high school start times. J. Clin. Sleep Med. 7, U145-U181.
148. Waage, S., Moen, B. E., Pallesen, S., Eriksen, H. R., Ursin, H., Åkerstedt, T., and Bjorvatn, B. (2009). Shift work disorder among oil rig workers in the North Sea. Sleep 32, 558-565.
149. Wahlstrom, K. (2010). School start time and sleepy teens. Arch. Pediatr. Adolesc. Med. 164, 676-677.
150. Wakamatsu, H., Yoshinobu, Y., Aida, R., Moriya, T., Akiyama, M., and Shibata, S. (2001). Restricted-feeding-induced anticipatory activity rhythm is associated with a phase-shift of the expression of mPer1 and mPer2 mRNA in the cerebral cortex and hippocampus but not in the suprachiasmatic nucleus of mice. Eur. J. Neurosci. 13, 1190-1196.
151. Wang, L. M., Dragich, J. M., Kudo, T., Odom, I. H., Welsh, D. K., O'Dell, T. J., and Colwell, C. S. (2009). Expression of the circadian clock gene Period2 in the hippocampus: possible implications for synaptic plasticity and learned behaviour. ASNNeuro. 1. Pii: e00012.
152. Wertz, A. T., Ronda, J. M., Czeisler, C. A., and Wright, K. P. Jr. (2006). Effects of sleep inertia on cognition. JAMA295, 163-164.
153. Wolfson, A. R., Spaulding, N. L., Dandrow, C., and Baroni, E. M. (2007). Middle school start times: the importance of a good night's sleep for young adolescents. Behav. SleepMed. 5, 194-209.
154. Wright, K. P. Jr., Badia, P., and Wauquier, A. (1995). Topographical and temporal patterns of brain activity during the transition from wakefulness to sleep. Sleep 18, 880-889.
155. Wright, K. P. Jr., Drake, C. L., and Lockley, S. W. (2008). "Diagnostic tools for circadian rhythm sleep disorders" in Handbook of Sleep Disorders, ed C. A. Kushida (New York, NY: Taylor and Francis Group), 147-175.
156. Wright, K. P. Jr., Gronfier, C., Duffy, J. E., and Czeisler, C. A. (2005). Intrinsic period and light intensity determine the phase relationship between melatonin and sleep in humans. J. Biol. Rhythms 20, 168-177.
157. Wright, K. P. Jr., Hughes, R. J., Kronauer, R. E., Dijk, D. J., and Czeisler, C. A. (2001). Intrinsic near-24-h pacemaker period determines limits of circadian entrainment to a weak synchronizer in humans. Proc. Natl. Acad. Sci. U.S.A. 98, 14027-14032.
158. Wright, K. P. Jr., Hull, J. T., and Czeisler, C. A. (2002). Relationship between alertness, performance, and body temperature in humans. Am. J. Physiol. 283, R1370-R1377.
159. Wright, K. P. Jr., Hull, J. T., Hughes, R. J., Ronda, J. M., and Czeisler, C. A. (2006). Sleep and wakefulness out of phase with internal biological time impairs learning in humans. J. Cogn. Neurosci. 18, 508-521.
160. Wyatt, J. K., Cajochen, C., Ritz-De Cecco, A., Czeisler, C. A., and
161. Wyatt, J. K., Dijk, D. J., Ritz-De Cecco, A., Ronda, J. M., and Czeisler, C. A. (2006). Sleep-facilitating effect of exogenous melatonin in healthy young men and women is circadian-phase dependent. Sleep 29, 609-618.
162. Yamazaki, S., Numano, R., Abe, M., Hida, A., Takahashi, R., Ueda, M., Block, G. D., Sakaki, Y., Menaker, M., and Tei, H. (2000). Resetting central and peripheral circadian oscillators in transgenic rats. Science 288, 682-685.
163. Yanai, K., and Tashiro, M. (2007). The physiological and pathophysiological roles of neuronal histamine: an insight from human positron emission tomography studies. Pharmacol. Ther. 113, 1-15.
164. Zhou, X., Ferguson, S. A., Matthews, R. W., Sargent, C., Darwent, D., Kennaway, D. J., and Roach, G. D. (2011a). Dynamics of neurobe-havioral performance variability under forced desynchrony: evidence of state instability. Sleep 34, 57-63.
165. Zhou, X., Ferguson, S. A., Matthews, R. W., Sargent, C., Darwent, D., Kennaway, D. J., and Roach, G. D. (2011b). Sleep, wake and phase dependent changes in neurobehav-ioral function under forced desynchrony. Sleep 34, 931-941.
166. Zhou, X., Ferguson, S. A., Matthews, R. W., Sargent, C., Darwent, D., Kennaway, D. J., and Roach, G. D. (2012). Mismatch between subjective alertness and objective performance under sleep restriction is greatest during the biological night. J. Sleep Res. 21, 40-9.