Effects of light on human circadian rhythms, sleep and mood

Christine Blume, Corrado Garbazza, Manuel Spitschan, Christine Blume, Corrado Garbazza, Manuel Spitschan

Abstract

Humans live in a 24-hour environment, in which light and darkness follow a diurnal pattern. Our circadian pacemaker, the suprachiasmatic nuclei (SCN) in the hypothalamus, is entrained to the 24-hour solar day via a pathway from the retina and synchronises our internal biological rhythms. Rhythmic variations in ambient illumination impact behaviours such as rest during sleep and activity during wakefulness as well as their underlying biological processes. Rather recently, the availability of artificial light has substantially changed the light environment, especially during evening and night hours. This may increase the risk of developing circadian rhythm sleep-wake disorders (CRSWD), which are often caused by a misalignment of endogenous circadian rhythms and external light-dark cycles. While the exact relationship between the availability of artificial light and CRSWD remains to be established, nocturnal light has been shown to alter circadian rhythms and sleep in humans. On the other hand, light can also be used as an effective and noninvasive therapeutic option with little to no side effects, to improve sleep,mood and general well-being. This article reviews our current state of knowledge regarding the effects of light on circadian rhythms, sleep, and mood.

Keywords: Artificial light; Circadian rhythms; Depression; Light therapy; Natural light.

Conflict of interest statement

Conflict of interest C. Blume, C. Garbazza and M. Spitschan declare that they have no relevant competing interests.

Figures

Fig. 1
Fig. 1
Input and output pathways to/from the suprachiasmatic nuclei (SCN). The photic input pathways that relay information about the intensity and spectral composition of ambient light are the retinohypothalamic tract (RHT) and the geniculohypothalamic tract (GHT), which connects retina and SCN via the intergeniculate leaflet (IGL) in the thalamus. Additionally, the SCN also receive non-photic information from the raphe nuclei (RN) via the raphe-hypothalamic tract (raphe-HT) and from the pineal gland. The main output is from the SCN to the serotonergic raphe nuclei (RN, receive information about the phase of the circadian clock and regulate vigilance state of the body) and the pineal gland, where melatonin is produced. Input and output pathways form reciprocal loops
Fig. 2
Fig. 2
Spectral power distributions of common light sources in our environment. a Spectral power distributions of daylights at different correlated colour temperatures (CCT; 4000 K; 6500 K; 10,000 K). Spectra are normalised to 555 nm. b Spectral power distributions of a white LED (top), a fluorescent source at 3000 K (middle), and an incandescent source (tungsten-filament; 2856 K, bottom). All three artificial sources have the same luminous flux (normalised to 100 lm), and approximately the same colour temperature (2700–3000 K), but the spectra are very different in shape and scale (see y axis)
Fig. 3
Fig. 3
Overview of the retina photoreceptors. a Schematic view of the eye with the retina at the back of the eye (the fundus), containing cones, rods and the intrinsically photosensitive retinal ganglion cells (ipRGCs) expressing the photopigment melanopsin. b Spectral sensitivities of the photoreceptors in the human eye

References

    1. Achermann P, Dijk D-J, Brunner DP, et al. A model of human sleep homeostasis based on EEG slow-wave activity: Quantitative comparison of data and simulations. Brain Res Bull. 1993;31:97–113. doi: 10.1016/0361-9230(93)90016-5.
    1. Al-Karawi D, Jubair L. Bright light therapy for nonseasonal depression: meta-analysis of clinical trials. J Affect Disord. 2016;198:64–71. doi: 10.1016/j.jad.2016.03.016.
    1. Allen AE, Hazelhoff EM, Martial FP, et al. Exploiting metamerism to regulate the impact of a visual display on alertness and melatonin suppression independent of visual appearance. Sleep. 2018 doi: 10.1093/sleep/zsy100.
    1. Bailes HJ, Lucas RJ. Human melanopsin forms a pigment maximally sensitive to blue light (lambdamax approximately 479 nm) supporting activation of G(q/11) and G(i/o) signalling cascades. Proc Biol Sci. 2013;280:20122987. doi: 10.1098/rspb.2012.2987.
    1. Beauchamp MT, Lundgren JD. A systematic review of bright light therapy for eating disorders. Prim Care Companion CNS Disord. 2016 doi: 10.4088/PCC.16r02008.
    1. Beauchemin KM, Hays P. Sunny hospital rooms expedite recovery from severe and refractory depressions. J Affect Disord. 1996;40:49–51. doi: 10.1016/0165-0327(96)00040-7.
    1. Benedetti F, Barbini B, Fulgosi MC, et al. Combined total sleep deprivation and light therapy in the treatment of drug-resistant bipolar depression: acute response and long-term remission rates. J Clin Psychiatry. 2005;66:1535–1540. doi: 10.4088/JCP.v66n1207.
    1. Benedetti F, Colombo C, Barbini B, et al. Morning sunlight reduces length of hospitalization in bipolar depression. J Affect Disord. 2001;62:221–223. doi: 10.1016/S0165-0327(00)00149-X.
    1. Blume C, Angerer M, Raml M, et al. Healthier rhythm, healthier brain? Integrity of circadian melatonin and temperature rhythms relates to the clinical state of brain-injured patients. Eur J Neurol. 2019 doi: 10.1111/ene.13935.
    1. Blume C, Lechinger J, Santhi N, et al. Significance of circadian rhythms in severely brain-injured patients: a clue to consciousness? Baillieres Clin Neurol. 2017 doi: 10.1212/WNL.0000000000003942.
    1. Boivin DB, Czeisler CA, Dijk D-J, et al. Complex interaction of the sleep-wake cycle and circadian phase modulates mood in healthy subjects. Arch Gen Psychiatry. 1997;54:145–152. doi: 10.1001/archpsyc.1997.01830140055010.
    1. Bollinger T, Schibler U. Circadian rhythms—from genes to physiology and disease. Swiss Med Wkly. 2014 doi: 10.4414/smw.2014.13984.
    1. Borbély AA. A two process model of sleep regulation. Hum Neurobiol. 1982;1(3):195–204.
    1. Borbély AA, Achermann P, Trachsel L, et al. Sleep initiation and initial sleep intensity: interactions of homeostatic and circadian mechanisms. J Biol Rhythms. 1989;4:37–48. doi: 10.1177/074873048900400205.
    1. Borbély AA, Daan S, Wirz-Justice A, et al. The two-process model of sleep regulation: a reappraisal. J Sleep Res. 2016;25:131–143. doi: 10.1111/jsr.12371.
    1. Boubekri M, Cheung IN, Reid KJ, et al. Impact of windows and daylight exposure on overall health and sleep quality of office workers: a case-control pilot study. J Clin Sleep Med. 2014;10:603–611.
    1. Brainard GC, Hanifin JP, Greeson JM, et al. Action spectrum for melatonin regulation in humans: evidence for a novel circadian photoreceptor. J Neurosci. 2001;21:6405–6412. doi: 10.1523/JNEUROSCI.21-16-06405.2001.
    1. Bromundt V, Köster M, Georgiev-Kill A, et al. Sleep–wake cycles and cognitive functioning in schizophrenia. Br J Psychiatry. 2011;198:269–276. doi: 10.1192/bjp.bp.110.078022.
    1. Bromundt V, Wirz-Justice A, Kyburz S, et al. Circadian sleep-wake cycles, well-being, and light therapy in borderline personality disorder. J Pers Disord. 2013;27:680–696. doi: 10.1521/pedi_2012_26_057.
    1. Brown TM, Tsujimura S, Allen AE, et al. Melanopsin-based brightness discrimination in mice and humans. Curr Biol. 2012;22:1134–1141. doi: 10.1016/j.cub.2012.04.039.
    1. Burkhalter H, Wirz‐Justice A, Denhaerynck K, et al. The effect of bright light therapy on sleep and circadian rhythms in renal transplant recipients: a pilot randomized, multicentre wait‐list controlled trial. Transpl Int. 2015;28:59–70. doi: 10.1111/tri.12443.
    1. Cajochen C. Chronobiologie: Licht-und Wachtherapie bei psychiatrischen Erkrankungen. Psych Up2date. 2013;7:173–184. doi: 10.1055/s-0033-1343181.
    1. Cajochen C, Kräuchi K, Wirz‐Justice A. Role of melatonin in the regulation of human circadian rhythms and sleep. J Neuroendocrinol. 2003;15:432–437. doi: 10.1046/j.1365-2826.2003.00989.x.
    1. Cajochen C, Munch M, Kobialka S, et al. High sensitivity of human melatonin, alertness, thermoregulation, and heart rate to short wavelength light. J Clin Endocrinol Metab. 2005;90:1311–1316. doi: 10.1210/jc.2004-0957.
    1. Cao D, Chang A, Gai S. Evidence for an impact of melanopsin activation on unique white perception. J Opt Soc Am A Opt Image Sci Vis. 2018;35:B287–B291. doi: 10.1364/JOSAA.35.00B287.
    1. Chang A-M, Aeschbach D, Duffy JF, et al. Evening use of light-emitting eReaders negatively affects sleep, circadian timing, and next-morning alertness. Proc. Natl. Acad. Sci. U.S.A. 2015;112:1232–1237. doi: 10.1073/pnas.1418490112.
    1. Chang AM, Scheer FA, Czeisler CA. The human circadian system adapts to prior photic history. J Physiol. 2011;589:1095–1102. doi: 10.1113/jphysiol.2010.201194.
    1. Chellappa SL, Steiner R, Oelhafen P, et al. Acute exposure to evening blue‐enriched light impacts on human sleep. J Sleep Res. 2013;22:573–580. doi: 10.1111/jsr.12050.
    1. Christensen MA, Bettencourt L, Kaye L, et al. Direct measurements of smartphone screen-time: relationships with demographics and sleep. PLoS ONE. 2016;11:e0165331. doi: 10.1371/journal.pone.0165331.
    1. Ciarleglio C, Resuehr H, Mcmahon D. Interactions of the serotonin and circadian systems: nature and nurture in rhythms and blues. Neuroscience. 2011;197:8–16. doi: 10.1016/j.neuroscience.2011.09.036.
    1. Cie . CIE system for metrology of optical radiation for ipRGC-influenced responses to light. Vienna, Austria: CIE Central Bureau; 2018.
    1. Dacey DM, Liao HW, Peterson BB, et al. Melanopsin-expressing ganglion cells in primate retina signal colour and irradiance and project to the LGN. Nature. 2005;433:749–754. doi: 10.1038/nature03387.
    1. De Rui M, Middleton B, Sticca A, et al. Sleep and circadian rhythms in hospitalized patients with decompensated cirrhosis: effect of light therapy. Neurochem Res. 2015;40:284–292. doi: 10.1007/s11064-014-1414-z.
    1. Dijk D-J, Czeisler CA. 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. 1995;15:3526–3538. doi: 10.1523/JNEUROSCI.15-05-03526.1995.
    1. Dijk D-J, Czeisler CA. Paradoxical timing of the circadian rhythm of sleep propensity serves to consolidate sleep and wakefulness in humans. Neurosci Lett. 1994;166:63–68. doi: 10.1016/0304-3940(94)90841-9.
    1. Engwall M, Fridh I, Johansson L, et al. Lighting, sleep and circadian rhythm: an intervention study in the intensive care unit. Intensive Crit Care Nurs. 2015;31:325–335. doi: 10.1016/j.iccn.2015.07.001.
    1. European Union (2011) European Lighting Standard. In: EN12464–1:2011.
    1. Fernandez DC, Fogerson PM, Lazzerini Ospri L, et al. Light affects mood and learning through distinct retina-brain pathways. Cell. 2018;175:71–84.e18. doi: 10.1016/j.cell.2018.08.004.
    1. Figueiro MG, Steverson B, Heerwagen J, et al. The impact of daytime light exposures on sleep and mood in office workers. Sleep Health. 2017;3:204–215. doi: 10.1016/j.sleh.2017.03.005.
    1. Gallin PF, Terman M, Reme CE, et al. Ophthalmologic examination of patients with seasonal affective disorder, before and after bright light therapy. Am J Ophthalmol. 1995;119:202–210. doi: 10.1016/S0002-9394(14)73874-7.
    1. Goel N, Terman M, Terman JS, et al. Controlled trial of bright light and negative air ions for chronic depression. Psychol Med. 2005;35:945–955. doi: 10.1017/S0033291705005027.
    1. Gooley JJ, Rajaratnam SM, Brainard GC, et al. Spectral responses of the human circadian system depend on the irradiance and duration of exposure to light. Sci Transl Med. 2010;2:31ra33. doi: 10.1126/scitranslmed.3000741.
    1. Gronfier C, Wright KP, Jr, Kronauer RE, et al. Efficacy of a single sequence of intermittent bright light pulses for delaying circadian phase in humans. Am. J. Physiol. Endocrinol. Metab. 2004 doi: 10.1152/ajpendo.00385.2003.
    1. Hebert M, Martin SK, Lee C, et al. The effects of prior light history on the suppression of melatonin by light in humans. J Pineal Res. 2002;33:198–203. doi: 10.1034/j.1600-079X.2002.01885.x.
    1. Ibuka N, Kawamura H. Loss of circadian rhythm in sleep-wakefulness cycle in the rat by suprachiasmatic nucleus lesions. Brain Res Brain Res Protoc. 1975;96:76–81.
    1. Kaiser C, Kaufmann C, Leutritz T, et al. The human habenula is responsive to changes in luminance and circadian rhythm. Neuroimage. 2019;189:581–588. doi: 10.1016/j.neuroimage.2019.01.064.
    1. Krasnik C, Montori VM, Guyatt GH, et al. The effect of bright light therapy on depression associated with premenstrual dysphoric disorder. Am J Obstet Gynecol. 2005;193:658–661. doi: 10.1016/j.ajog.2005.01.055.
    1. Kronauer RE, Forger DB, Jewett ME. Quantifying human circadian pacemaker response to brief, extended, and repeated light stimuli over the phototopic range. J Biol Rhythms. 1999;14:501–516. doi: 10.1177/074873049901400609.
    1. Lam RW, Goldner EM, Solyom L, et al. A controlled study of light therapy for bulimia nervosa. Am J Psychiatry. 1994;151:744. doi: 10.1176/ajp.151.5.744.
    1. Lemola S, Perkinson-Gloor N, Brand S, et al. Adolescents’ electronic media use at night, sleep disturbance, and depressive symptoms in the smartphone age. J Youth Adolesc. 2015;44:405–418. doi: 10.1007/s10964-014-0176-x.
    1. Lewy AJ, Wehr TA, Goodwin FK, et al. Light suppresses melatonin secretion in humans. Science. 1980;210:1267–1269. doi: 10.1126/science.7434030.
    1. Lieverse R, Van Someren EJ, Nielen MM, et al. Bright light treatment in elderly patients with nonseasonal major depressive disorder: a randomized placebo-controlled trial. Arch Gen Psychiatry. 2011;68:61–70. doi: 10.1001/archgenpsychiatry.2010.183.
    1. Martiny K. Adjunctive bright light in non‐seasonal major depression. Acta Psychiatr Scand. 2004;110:7–28. doi: 10.1111/j.1600-0447.2004.00460_2.x.
    1. Meyer-Bernstein E, Morin L. Differential serotonergic innervation of the suprachiasmatic nucleus and the intergeniculate leaflet and its role in circadian rhythm modulation. J Neurosci. 1996;16:2097–2111. doi: 10.1523/JNEUROSCI.16-06-02097.1996.
    1. Mohawk JA, Green CB, Takahashi JS. Central and peripheral circadian clocks in mammals. Annu Rev Neurosci. 2012;35:445–462. doi: 10.1146/annurev-neuro-060909-153128.
    1. Moore RY. Suprachiasmatic nucleus in sleep–wake regulation. Sleep Med. 2007;8:27–33. doi: 10.1016/j.sleep.2007.10.003.
    1. Mouret J, Coindet J, Debilly G, et al. Suprachiasmatic nuclei lessions in the rat: alterations in sleep circadian rhythms. Electroencephalogr Clin Neurophysiol. 1978;45:402–408. doi: 10.1016/0013-4694(78)90191-8.
    1. Münch M, Kobialka S, Steiner R, et al. Wavelength-dependent effects of evening light exposure on sleep architecture and sleep EEG power density in men. Am. J. Physiol. Regul. Integr. Comp. Physiol. 2006;290:R1421–R1428. doi: 10.1152/ajpregu.00478.2005.
    1. Najjar RP, Zeitzer JM. Temporal integration of light flashes by the human circadian system. J Clin Invest. 2016;126:938–947. doi: 10.1172/JCI82306.
    1. Nowozin C, Wahnschaffe A, Rodenbeck A, et al. Applying Melanopic lux to measure biological light effects on Melatonin suppression and subjective sleepiness. Curr Alzheimer Res. 2017;14:1042–1052. doi: 10.2174/1567205014666170523094526.
    1. Partonen T, Pandi-Perumal S. Seasonal affective disorder: practice and research. Oxford, UK: Oxford University Press; 2010.
    1. Provencio I, Rodriguez IR, Jiang G, et al. A novel human opsin in the inner retina. J Neurosci. 2000;20:600–605. doi: 10.1523/JNEUROSCI.20-02-00600.2000.
    1. Rahman SA, Hilaire StMA, Gronfier C, et al. Functional decoupling of melatonin suppression and circadian phase resetting in humans. J Physiol. 2018;596:2147–2157. doi: 10.1113/JP275501.
    1. Rångtell FH, Ekstrand E, Rapp L, et al. Two hours of evening reading on a self-luminous tablet vs. reading a physical book does not alter sleep after daytime bright light exposure. Sleep Med. 2016;23:111–118. doi: 10.1016/j.sleep.2016.06.016.
    1. Riemersma-Van Der Lek RF, Swaab DF, Twisk J, et al. Effect of bright light and melatonin on cognitive and noncognitive function in elderly residents of group care facilities: a randomized controlled trial. JAMA. 2008;299:2642–2655. doi: 10.1001/jama.299.22.2642.
    1. Rimmer DW, Boivin DB, Shanahan TL, et al. Dynamic resetting of the human circadian pacemaker by intermittent bright light. Am. J. Physiol. Regul. Integr. Comp. Physiol. 2000;279:R1574–R1579. doi: 10.1152/ajpregu.2000.279.5.R1574.
    1. Roenneberg T, Wirz-Justice A, Merrow M. Life between clocks: daily temporal patterns of human Chronotypes. J Biol Rhythms. 2003;18:80–90. doi: 10.1177/0748730402239679.
    1. Rosenthal NE, Sack DA, Gillin JC, et al. Seasonal affective disorder: a description of the syndrome and preliminary findings with light therapy. Arch Gen Psychiatry. 1984;41:72–80. doi: 10.1001/archpsyc.1984.01790120076010.
    1. Rybak YE, Mcneely HE, Mackenzie BE, et al. An open trial of light therapy in adult attention-deficit/hyperactivity disorder. J Clin Psychiatry. 2006;67:1527–1535. doi: 10.4088/JCP.v67n1006.
    1. Sack RL, Auckley D, Auger RR, et al. Circadian rhythm sleep disorders: part I, basic principles, shift work and jet lag disorders. Sleep. 2007;30:1460–1483. doi: 10.1093/sleep/30.11.1460.
    1. Sack RL, Lewy AJ, Hughes RJ. Use of melatonin for sleep and circadian rhythm disorders. Ann Med. 1998;30:115–121. doi: 10.3109/07853899808999393.
    1. Santhi N, Thorne HC, Van Der Veen DR, et al. The spectral composition of evening light and individual differences in the suppression of melatonin and delay of sleep in humans. J Pineal Res. 2012;53:47–59. doi: 10.1111/j.1600-079X.2011.00970.x.
    1. Schwartz RS, Olds J. The psychiatry of light. Harv Rev Psychiatry. 2015;23:188–194. doi: 10.1097/HRP.0000000000000078.
    1. Schweizer A, Berchtold A, Barrense-Dias Y, et al. Adolescents with a smartphone sleep less than their peers. Eur J Pediatr. 2017;176:131–136. doi: 10.1007/s00431-016-2823-6.
    1. Sheaves B, Freeman D, Isham L, et al. Stabilising sleep for patients admitted at acute crisis to a psychiatric hospital (OWLS): an assessor-blind pilot randomised controlled trial. Psychol Med. 2018;48:1694–1704. doi: 10.1017/S0033291717003191.
    1. Shochat T, Martin J, Marler M, et al. Illumination levels in nursing home patients: effects on sleep and activity rhythms. J Sleep Res. 2000;9:373–379. doi: 10.1046/j.1365-2869.2000.00221.x.
    1. Smith KA, Schoen MW, Czeisler CA. Adaptation of human pineal melatonin suppression by recent photic history. J Clin Endocrinol Metab. 2004;89:3610–3614. doi: 10.1210/jc.2003-032100.
    1. Souman JL, Borra T, de Goijer I, et al. Spectral tuning of white light allows for strong reduction in melatonin suppression without changing illumination level or color temperature. J Biol Rhythms. 2018;33(4):420–431. doi: 10.1177/0748730418784041.
    1. Spitschan M. Melanopsin contributions to non-visual and visual function. Curr Opini Behav Sci. 2019;30:67–72. doi: 10.1016/j.cobeha.2019.06.004.
    1. Spitschan M, Jain S, Brainard DH, et al. Opponent melanopsin and S-cone signals in the human pupillary light response. Proc Natl Acad Sci U S A. 2014;111:15568–15572. doi: 10.1073/pnas.1400942111.
    1. Spitschan M, Aguirre GK, Brainard DH, et al. Variation of outdoor illumination as a function of solar elevation and light pollution. Sci Rep. 2016;6:26756. doi: 10.1038/srep26756.
    1. Spitschan M, Bock AS, Ryan J, et al. The human visual cortex response to melanopsin-directed stimulation is accompanied by a distinct perceptual experience. Proc. Natl. Acad. Sci. U. S. A. 2017;114:12291–12296. doi: 10.1073/pnas.1711522114.
    1. Spitschan M, Stefani O, Blattner P, et al. How to report light exposure in human chronobiology and sleep research experiments. Clocks. Sleep. 2019;1(3):280–289.
    1. Stephan FK, Nunez AA. Elimination of circadian rhythms in drinking, activity, sleep, and temperature by isolation of the suprachiasmatic nuclei. Behav Biol. 1977;20:1–16. doi: 10.1016/S0091-6773(77)90397-2.
    1. Stothard ER, Mchill AW, Depner CM, et al. Circadian entrainment to the natural light-dark cycle across seasons and the weekend. Curr Biol. 2017;27:508–513. doi: 10.1016/j.cub.2016.12.041.
    1. Terman JS, Terman M, Lo E-S, et al. Circadian time of morning light administration and therapeutic response in winter depression. Arch Gen Psychiatry. 2001;58:69–75. doi: 10.1001/archpsyc.58.1.69.
    1. Terman M, Terman JS, Ross DC. A controlled trial of timed bright light and negative air ionization for treatment of winter depression. Arch Gen Psychiatry. 1998;55:875–882. doi: 10.1001/archpsyc.55.10.875.
    1. Thapan K, Arendt J, Skene DJ. An action spectrum for melatonin suppression: evidence for a novel non-rod, non-cone photoreceptor system in humans. J Physiol. 2001;535:261–267. doi: 10.1111/j.1469-7793.2001.t01-1-00261.x.
    1. Thorpy M. International classification of sleep disorders. In: Chokroverty S, editor. Sleep disorders medicine: basic science, technical considerations and clinical aspects. New York, NY: Springer; 2017. pp. 475–484.
    1. Videnovic A, Klerman EB, Wang W, et al. Timed light therapy for sleep and daytime sleepiness associated with parkinson disease: a randomized clinical trial. Jama Neurol. 2017;74:411–418. doi: 10.1001/jamaneurol.2016.5192.
    1. Viola AU, James LM, Schlangen LJ, et al. Blue-enriched white light in the workplace improves self-reported alertness, performance and sleep quality. Scand J Work Environ Health. 2008;34(4):297–306. doi: 10.5271/sjweh.1268.
    1. Wams EJ, Woelders T, Marring I, et al. Linking light exposure and subsequent sleep: a field Polysomnography study in humans. Sleep. 2017 doi: 10.1093/sleep/zsx165.
    1. Webler F, Spitschan M, Foster R, et al. What is the “spectral diet” of humans? Curr Opini Behav Sci. 2019;30:80–86. doi: 10.1016/j.cobeha.2019.06.006.
    1. Wehr TA. The durations of human melatonin secretion and sleep respond to changes in daylength (photoperiod) J Clin Endocrinol Metab. 1991;73:1276–1280. doi: 10.1210/jcem-73-6-1276.
    1. Wehr TA, Moul DE, Barbato G, et al. Conservation of photoperiod-responsive mechanisms in humans. Am. J. Physiol. Regul. Integr. Comp. Physiol. 1993;265:R846–R857. doi: 10.1152/ajpregu.1993.265.4.R846.
    1. Wirz-Justice A, Bader A, Frisch U, et al. A randomized, double-blind, placebo-controlled study of light therapy for antepartum depression. J Clin Psychiatry. 2011;72:986–993. doi: 10.4088/JCP.10m06188blu.
    1. Wirz-Justice A, Benedetti F, Terman M. Chronotherapeutics for affective disorders: a clinician’s manual for light and wake therapy. 2. Basel, Switzerland: Karger Medical and Scientific Publishers; 2013.
    1. Wirz-Justice A, Bromundt V. Lichttherapie. Schlaf. 2013;2:20–29. doi: 10.1055/s-0038-1626044.
    1. Wirz-Justice A, Graw P, Kräuchi K, et al. ‘Natural’light treatment of seasonal affective disorder. J Affect Disord. 1996;37:109–120. doi: 10.1016/0165-0327(95)00081-X.
    1. Woelders T, Leenheers T, Gordijn MCM, et al. Melanopsin- and L-cone-induced pupil constriction is inhibited by S- and M-cones in humans. Proc Natl Acad Sci U S A. 2018;115:792–797. doi: 10.1073/pnas.1716281115.
    1. Wright HR, Lack LC. Effect of light wavelength on suppression and phase delay of the melatonin rhythm. Chronobiol Int. 2001;18:801–808. doi: 10.1081/CBI-100107515.
    1. Wright KP, Mchill AW, Birks BR, et al. Entrainment of the human circadian clock to the natural light-dark cycle. Curr Biol. 2013;23:1554–1558. doi: 10.1016/j.cub.2013.06.039.
    1. Wulff K, Gatti S, Wettstein JG, et al. Sleep and circadian rhythm disruption in psychiatric and neurodegenerative disease. Nat Rev Neurosci. 2010;11:589. doi: 10.1038/nrn2868.
    1. Yetish G, et al. Natural sleep and its seasonal variations in three pre-industrial societies. Current Biology. 2015;25(21):2862–2868. doi: 10.1016/j.cub.2015.09.046.
    1. Youngstedt SD, Elliott JA, Kripke DF. Human circadian phase—response curves for exercise. J. Physiol. 2019;597:2253–2268. doi: 10.1113/JP276943.
    1. Zeitzer JM. When is a proxy not a proxy? The foibles of studying non-image forming light. J. Physiol. 2018;596:2029–2030. doi: 10.1113/JP276076.
    1. Zeitzer JM, Dijk D-J, Kronauer RE, et al. Sensitivity of the human circadian pacemaker to nocturnal light: melatonin phase resetting and suppression. J. Physiol. 2000;526:695–702. doi: 10.1111/j.1469-7793.2000.00695.x.
    1. Zeitzer JM, Fisicaro RA, Ruby NF, et al. Millisecond flashes of light phase delay the human circadian clock during sleep. J Biol Rhythms. 2014;29:370–376. doi: 10.1177/0748730414546532.
    1. Zele AJ, Adhikari P, Feigl B, et al. Cone and melanopsin contributions to human brightness estimation. J Opt Soc Am A Opt Image Sci Vis. 2018;35:B19–B25. doi: 10.1364/JOSAA.35.000B19.
    1. Zhang B-B, Yao Y-Y, Zhang H-F, et al. Left Habenula mediates light-preference behavior in Zebrafish via an asymmetrical visual pathway. Neuron. 2017;93:914–928.e914. doi: 10.1016/j.neuron.2017.01.011.

Source: PubMed

3
구독하다