非成像视觉的光信号转导与环路结构功能
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摘要
感知外界环境并做出响应是生物体的基本特征,而感光是多数生物最重要的感知觉之一.生命体为此特化出了复杂的感光机制,不仅用于形成图像视觉,也帮助生物根据光环境调节相应的生理功能,如瞳孔光反射和生物节律的光授时.这些感光功能统称为非成像视觉功能,主要由一类新近发现的自感光视网膜神经节细胞所介导.自感光视网膜神经节细胞有别于传统的视锥视杆细胞,自发现以来,逐渐积累的研究证实了这类神经元投射到丰富的皮层下结构,并参与包括睡眠节律和情绪调节等多种生理功能.本文旨在回顾有关自感光视网膜神经节细胞光信号转导与皮层下环路的重要发现和最新进展,并针对领域的趋势和待解决问题进行展望.
Perception of light not only results in image vision,but also modulates a number of biological functions that do not involve image formation,such as pupil light reflex and photoentrainment of circadian rhythm.Such non-image-forming vision is primarily mediated by a subset of retinal ganglion cells,termed as intrinsically photosensitive retinal ganglion cells(ipRGCs),which express the photopigment melanopsin thereby exhibiting intrinsic photosensitivity.Since the groundbreaking discovery of ipRGCs approximately two decades ago,ample evidence suggests that ipRGCs project to various subcortical nuclei,and probably participate in a wider range of physiological processes than initially known.This article reviews important laboratory and clinical evidence and recent progress regarding phototransduction of ipRGCs and brain circuitry in non-image vision.
Perception of light not only results in image vision,but also modulates a number of biological functions that do not involve image formation,such as pupil light reflex and photoentrainment of circadian rhythm.Such non-image-forming vision is primarily mediated by a subset of retinal ganglion cells,termed as intrinsically photosensitive retinal ganglion cells(ipRGCs),which express the photopigment melanopsin thereby exhibiting intrinsic photosensitivity.Since the groundbreaking discovery of ipRGCs approximately two decades ago,ample evidence suggests that ipRGCs project to various subcortical nuclei,and probably participate in a wider range of physiological processes than initially known.This article reviews important laboratory and clinical evidence and recent progress regarding phototransduction of ipRGCs and brain circuitry in non-image vision.
引文
1 Provencio I,Jiang G,De Grip W J,et al.Melanopsin:an opsin in melanophores,brain,and eye.Proc Natl Acad Sci USA,1998,95:340-345
2 Gooley J J,Lu J,Chou T C,et al.Melanopsin in cells of origin of the retinohypothalamic tract.Nat Neurosci,2001,4:1165-1165
3 Hattar S,Liao H W,Takao M,et al.Melanopsin-containing retinal ganglion cells:architecture,projections,and intrinsic photosensitivity.Science,2002,295:1065-1070
4 Provencio I,Rodriguez I R,Jiang G,et al.A novel human opsin in the inner retina.J Neurosci,2000,20:600-605
5 Lucas R J,Hattar S,Takao M,et al.Diminished pupillary light reflex at high irradiances in melanopsin-knockout mice.Science,2003,299:245-247
6 Freedman M S,Lucas R J,Soni B,et al.Regulation of mammalian circadian behavior by non-rod,non-cone,ocular photoreceptors.Science,1999,284:502-504
7 LeGates T A,Altimus C M,Wang H,et al.Aberrant light directly impairs mood and learning through melanopsin-expressing neurons.Nature,2012,491:594-598
8 Yau K W,Hardie R C.Phototransduction motifs and variations.Cell,2009,139:246-264
9 Arendt D.Evolution of eyes and photoreceptor cell types.Int J Dev Biol,2003,47:563-571
10 Lamb T D,Collin S P,PughE N.Evolution of the vertebrate eye:opsins,photoreceptors,retina and eye cup.Nat Rev Neurosci,2007,8:960-976
11 Terakita A.The opsins.Genome Biol,2005,6:213
12 Finn J T,Solessio E C,Yau K W.A cGMP-gated cation channel in depolarizing photoreceptors of the lizard parietal eye.Nature,1997,385:815-819
13 Xiong W H,Solessio E C,Yau K W.An unusual cGMP pathway underlying depolarizing light response of the vertebrate parietal-eye photoreceptor.NatNeurosci,1998,1:359-365
14 Xue T,Do M T H,Riccio A,et al.Melanopsin signalling in mammalian iris and retina.Nature,2011,479:67-73
15 Katz B,Minke B.Drosophila photoreceptors and signaling mechanisms.Front Cell Neurosci,2009,3:2
16 Hardie R C,Raghu P.Visual transduction in Drosophila.Nature,2001,413:186-193
17 Do M T H,Yau K W.Intrinsically photosensitive retinal ganglion cells.Physiological Rev,2010,90:1547-1581
18 Fain G L,Hardie R,Laughlin S B.Phototransduction and the evolution of photoreceptors.Curr Biol,2010,20:R114-R124
19 Del Pilar Gomez M,Angueyra J M,Nasi E.Light-transduction in melanopsin-expressing photoreceptors of Amphioxus.Proc Natl Acad Sci USA,2009,106:9081-9086
20 Mure L S,Rieux C,Hattar S,et al.Melanopsin-dependent nonvisual responses:evidence for photopigment bistability in vivo.J Biol Rhythms,2007,22:411-424
21 Walker M T,Brown R L,Cronin T W,et al.Photochemistry of retinal chromophore in mouse melanopsin.Proc Natl Acad Sci USA,2008,105:8861-8865
22 Emanuel A J,Do M TH.Melanopsin tristability for sustained and broadband phototransduction.Neuron,2015,85:1043-1055
23 Tu D C,Owens L A,Anderson L,et al.Inner retinal photoreception independent of the visual retinoid cycle.Proc Natl Acad Sci USA,2006,103:10426-10431
24 Doyle S E,Castrucci A M,McCall M,et al.Nonvisual light responses in the Rpe65 knockout mouse:rod loss restores sensitivity to the melanopsin system.Proc Natl Acad Sci USA,2006,103:10432-10437
25 Viney T J,Balint K,Hillier D,et al.Local retinal circuits of melanopsin-containing ganglion cells identified by transsynaptic viral tracing.Curr Biol,2007,17:981-988
26 Do M T H,Kang S H,Xue T,et al.Photon capture and signalling by melanopsin retinal ganglion cells.Nature,2009,457:281-287
27 Berson D M.Phototransduction in ganglion-cell photoreceptors.Pflugers Arch,2007,454:849-855
28 Dacey D M,Liao H W,Peterson B B,et al.Melanopsin-expressing ganglion cells in primate retina signal colour and irradiance and project to the LGN.Nature,2005,433:749-754
29 Graham D M,Wong K Y,Shapiro P,et al.Melanopsin ganglion cells use a membrane-associated rhabdomeric phototransduction cascade.J Neurophysiol,2008,99:2522-2532
30 Hartwick A T E,Bramley J R,Yu J,et al.Light-evoked calcium responses of isolated melanopsin-expressing retinal ganglion cells.J Neurosci,2007,27:13468-13480
31 Sekaran S,Lall G S,Ralphs K L,et al.2-aminoethoxydiphenylborane is an acute inhibitor of directly photosensitive retinal ganglion cell activity in vitro and in vivo.J Neurosci,2007,27:3981-3986
32 Dolph P J,Ranganathan R,Colley N J,et al.Arrestin function in inactivation of G protein-coupled receptor rhodopsin in vivo.Science,1993,260:1910-1916
33 Ranganathan R,Stevens CF.Arrestin binding determines the rate of inactivation of the G protein-coupled receptor rhodopsin in vivo.Cell,1995,81:841-848
34 Blasic J R,Lane Brown R,Robinson P R.Light-dependent phosphorylation of the carboxy tail of mouse melanopsin.Cell Mol Life Sci,2012,69:1551-1562
35 Cameron E G,Robinson P R.β-arrestin-dependent deactivation of mouse melanopsin.PLoS One,2014,9:ell3138
36 Godement P,Salailn J,Imbert M.Prenatal and postnatal development of retinogeniculate and retinocollicular projections in the mouse.J Comp Neurol,1984,230:552-575
37 Johnson R F,Morin L P,Moore R Y.Retinohypothalamic projections in the hamster and rat demonstrated using cholera toxin.Brain Res,1988,462:301-312
38 Pickard G E.The afferent connections of the suprachiasmatic nucleus of the golden hamster with emphasis on the retinohypothalamic projection.J Comp Neurol,1982,211:65-83
39 Moore R Y,Speh J C,Patrick Card J.The retinohypothalamic tract originates from a distinct subset of retinal ganglion cells.J Comp Neurol,1995,352:351-366
40 Berson D M,Dunn F A,Takao M.Phototransduction by retinal ganglion cells that set the circadian clock.Science,2002,295:1070-1073
41 Hattar S,Kumar M,Park A,et al.Central projections of melanopsin-expressing retinal ganglion cells in the mouse.J Comp Neurol,2006,497:326-349
42 Baver S B,Pickard G E,Sollars P J,et al.Two types of melanopsin retinal ganglion cell differentially innervate the hypothalamic suprachiasmatic nucleus and the olivary pretectal nucleus.Eur J Neurosci,2008,27:1763-1770
43 Hannibal J,Hindersson P,Knudsen S M,et al.The photopigment melanopsin is exclusively present in pituitary adenylate cyclase-activating polypeptide-containing retinal ganglion cells of the retinohypothalamic tract.J Neurosci,2002,22:RC191
44 Berson D.Strange vision:ganglion cells as circadian photoreceptors.Trends Neurosci,2003,26:314-320
45 Hirota T,Fukada Y.Resetting mechanism of central and peripheral circadian clocks in mammals.Zoolog Sci,2004,21:359-368
46 MorinL P,Blanchard J H,Provencio I.Retinal ganglion cell projections to the hamster suprachiasmatic nucleus,intergeniculate leaflet,and visual midbrain:bifurcation and melanopsin immunoreactivity.J Comp Neurol,2003,465:401-416
47 Harrington M E.The ventral lateral geniculate nucleus and the intergeniculate leaflet:interrelated structures in the visual and circadian systems.Neurosci Biobehav Rev,1997,21:705-727
48 Trejo L J,Cicerone C M.Cells in the pretectal olivary nucleus are in the pathway forthe direct light reflex of the pupil in the rat.Brain Res,1984,300:49-62
49 Gamlin P D R,Zhang H,Clarke R J.Luminance neurons in the pretectal olivary nucleus mediate the pupillary light reflex in the rhesus monkey.Exp Brain Res,1995,106:169
50 Young M J,Lund R D.The anatomical substrates subserving the pupillary light reflex in rats:origin of the consensual pupillary response.Neuroscience,1994,62:481-496
51 Muscat L,Huberman A D,Jordan C L,et al.Crossed and uncrossed retinal projections to the hamster circadian system.J Comp Neurol,2003,466:513-524
52 Hannibal J,Fahrenkrug J.Target areas innervated by PACAP-immunoreactive retinal ganglion cells.Cell Tissue Res,2004,316:99-113
53 Ecker J L,Dumitrescu O N,Wong K Y,et al.Melanopsin-expressing retinal ganglion-cell photoreceptors:cellular diversity and role in pattern vision.Neuron,2010,67:49-60
54 Hannibal J,Kankipati L,Strang C E,et al.Central projections of intrinsically photosensitive retinal ganglion cells in the macaque monkey.J Comp Neurol,2014,522:2231-2248
55 Hannibal J,Hindersson P,Ostergaard J,et al.Melanopsin is expressed in pacap-containing retinal ganglion cells of the human retinohypothalamic tract.Invest Ophthalmol Vis Sci,2004,45:4202-4209
56 Lucas R J,Douglas R H,Foster R G.Characterization of an ocular photopigment capable of driving pupillary constriction in mice.Nat Neurosci,2001,4:621-626
57 Semo M,Peirson S,Lupi D,et al.Melanopsin retinal ganglion cells and the maintenance of circadian and pupillary responses to light in aged rodless/coneless(rd/rd cl)mice.Eur J Neurosci,2003,17:1793-1801
58 Reppert S M,Weaver D R.Coordination of circadian timing in mammals.Nature,2002,418:935-941
59 Panda S,Sato T K,Castrucci A M,et al.Melanopsin(Opn4)requirement for normal light-induced circadian phase shifting.Science,2002,298:2213-2216
60 Roecklein K A,Rohan K J,Duncan WC,et al.A missense variant(P10L)of the melanopsin(OPN4)gene in seasonal affective disorder.J Affect Disord,2009,114:279-285
61 Abrahamson E E,Moore R Y.Suprachiasmatic nucleus in the mouse:retinal innervation,intrinsic organization and efferent projections.Brain Res,2001,916:172-191
62 Luo A H,Aston-Jones G.Circuit projection from suprachiasmatic nucleus to ventral tegmental area:a novel circadian output pathway.Eur J Neurosci,2009,29:748-760
63 Altimus C M,G(u|¨)ler A D,Villa K L,et al.Rods-cones and melanopsin detect light and dark to modulate sleep independent of image formation.Proc Natl Acad Sci USA,2008,105:19998-20003
64 Lupi D,Oster H,Thompson S,et al.The acute light-induction of sleep is mediated by OPN4-based photoreception.Nat Neurosci,2008,11:1068-1073
65 Kalsbeek A,Palm I F,La Fleur S E,et al.SCN outputs and the hypothalamic balance of life.J Biol Rhythms,2006,21:458-469
66 Czeisler C A,Shanahan T L,Klerman E B,et al.Suppression of melatonin secretion in some blind patients by exposure to bright light.N Engl J Med,1995,332:6-11
67 Srinivasan V,Smits M,Spence W,et al.Melatonin in mood disorders.World J Biol Psychiatry,2006,7:138-151
68 LeGates T A,Fernandez D C,Hattar S.Light as a central modulator of circadian rhythms,sleep and affect.Nat Rev Neurosci,2014,15:443-454
69 Perlis M L,Giles D E,Buysse D J,et al.Self-reported sleep disturbance as a prodromal symptom in recurrent depression.J Affect Disord,1997,42:209-212
70 Brown R E,Basheer R,McKenna J T,et al.Control of sleep and wakefulness.Physiol Rev,2012,92:1087-1187
71 Perrin F,Peigneux P,Fuchs S,et al.Nonvisual responses to light exposure in the human brain during the circadian night.Curr Biol,2004,14:1842-1846
72 Vandewalle G,Maquet P,Dijk D J.Light as a modulator of cognitive brain function.Trends Cogn Sci,2009,13:429-438
73 Vandewalle G,Collignon O,Hull J T,et al.Blue light stimulates cognitive brain activity in visually blind individuals.J Cogn Neurosci,2013,25:2072-2085
74 Vandewalle G,Gais S,Schabus M,et al.Wavelength-dependent modulation of brain responses to a working memory task by daytime light exposure.Cerebral Cortex,2007,17:2788-2795
75 Bonmati-Carrion M A,Arguelles-Prieto R,Martinez-Madrid M J,et al.Protecting the melatonin rhythm through circadian healthy light exposure.Int J Mol Sci,2014,15:23448-23500
76 Haus E L,Smolensky M H.Shift work and cancer risk:potential mechanistic roles of circadian disruption,light at night,and sleep deprivation.Sleep Med Rev,2013,17:273-284
77 Stephenson R,Lim J,Famina S,et al.Sleep-wake behavior in the rat.J Biol Rhythms,2012,27:490-501
78 Lewy A J,Lefler B J,Emens J S,et al.The circadian basis of winter depression.Proc Natl Acad Sci USA,2006,103:7414-7419
2 Gooley J J,Lu J,Chou T C,et al.Melanopsin in cells of origin of the retinohypothalamic tract.Nat Neurosci,2001,4:1165-1165
3 Hattar S,Liao H W,Takao M,et al.Melanopsin-containing retinal ganglion cells:architecture,projections,and intrinsic photosensitivity.Science,2002,295:1065-1070
4 Provencio I,Rodriguez I R,Jiang G,et al.A novel human opsin in the inner retina.J Neurosci,2000,20:600-605
5 Lucas R J,Hattar S,Takao M,et al.Diminished pupillary light reflex at high irradiances in melanopsin-knockout mice.Science,2003,299:245-247
6 Freedman M S,Lucas R J,Soni B,et al.Regulation of mammalian circadian behavior by non-rod,non-cone,ocular photoreceptors.Science,1999,284:502-504
7 LeGates T A,Altimus C M,Wang H,et al.Aberrant light directly impairs mood and learning through melanopsin-expressing neurons.Nature,2012,491:594-598
8 Yau K W,Hardie R C.Phototransduction motifs and variations.Cell,2009,139:246-264
9 Arendt D.Evolution of eyes and photoreceptor cell types.Int J Dev Biol,2003,47:563-571
10 Lamb T D,Collin S P,PughE N.Evolution of the vertebrate eye:opsins,photoreceptors,retina and eye cup.Nat Rev Neurosci,2007,8:960-976
11 Terakita A.The opsins.Genome Biol,2005,6:213
12 Finn J T,Solessio E C,Yau K W.A cGMP-gated cation channel in depolarizing photoreceptors of the lizard parietal eye.Nature,1997,385:815-819
13 Xiong W H,Solessio E C,Yau K W.An unusual cGMP pathway underlying depolarizing light response of the vertebrate parietal-eye photoreceptor.NatNeurosci,1998,1:359-365
14 Xue T,Do M T H,Riccio A,et al.Melanopsin signalling in mammalian iris and retina.Nature,2011,479:67-73
15 Katz B,Minke B.Drosophila photoreceptors and signaling mechanisms.Front Cell Neurosci,2009,3:2
16 Hardie R C,Raghu P.Visual transduction in Drosophila.Nature,2001,413:186-193
17 Do M T H,Yau K W.Intrinsically photosensitive retinal ganglion cells.Physiological Rev,2010,90:1547-1581
18 Fain G L,Hardie R,Laughlin S B.Phototransduction and the evolution of photoreceptors.Curr Biol,2010,20:R114-R124
19 Del Pilar Gomez M,Angueyra J M,Nasi E.Light-transduction in melanopsin-expressing photoreceptors of Amphioxus.Proc Natl Acad Sci USA,2009,106:9081-9086
20 Mure L S,Rieux C,Hattar S,et al.Melanopsin-dependent nonvisual responses:evidence for photopigment bistability in vivo.J Biol Rhythms,2007,22:411-424
21 Walker M T,Brown R L,Cronin T W,et al.Photochemistry of retinal chromophore in mouse melanopsin.Proc Natl Acad Sci USA,2008,105:8861-8865
22 Emanuel A J,Do M TH.Melanopsin tristability for sustained and broadband phototransduction.Neuron,2015,85:1043-1055
23 Tu D C,Owens L A,Anderson L,et al.Inner retinal photoreception independent of the visual retinoid cycle.Proc Natl Acad Sci USA,2006,103:10426-10431
24 Doyle S E,Castrucci A M,McCall M,et al.Nonvisual light responses in the Rpe65 knockout mouse:rod loss restores sensitivity to the melanopsin system.Proc Natl Acad Sci USA,2006,103:10432-10437
25 Viney T J,Balint K,Hillier D,et al.Local retinal circuits of melanopsin-containing ganglion cells identified by transsynaptic viral tracing.Curr Biol,2007,17:981-988
26 Do M T H,Kang S H,Xue T,et al.Photon capture and signalling by melanopsin retinal ganglion cells.Nature,2009,457:281-287
27 Berson D M.Phototransduction in ganglion-cell photoreceptors.Pflugers Arch,2007,454:849-855
28 Dacey D M,Liao H W,Peterson B B,et al.Melanopsin-expressing ganglion cells in primate retina signal colour and irradiance and project to the LGN.Nature,2005,433:749-754
29 Graham D M,Wong K Y,Shapiro P,et al.Melanopsin ganglion cells use a membrane-associated rhabdomeric phototransduction cascade.J Neurophysiol,2008,99:2522-2532
30 Hartwick A T E,Bramley J R,Yu J,et al.Light-evoked calcium responses of isolated melanopsin-expressing retinal ganglion cells.J Neurosci,2007,27:13468-13480
31 Sekaran S,Lall G S,Ralphs K L,et al.2-aminoethoxydiphenylborane is an acute inhibitor of directly photosensitive retinal ganglion cell activity in vitro and in vivo.J Neurosci,2007,27:3981-3986
32 Dolph P J,Ranganathan R,Colley N J,et al.Arrestin function in inactivation of G protein-coupled receptor rhodopsin in vivo.Science,1993,260:1910-1916
33 Ranganathan R,Stevens CF.Arrestin binding determines the rate of inactivation of the G protein-coupled receptor rhodopsin in vivo.Cell,1995,81:841-848
34 Blasic J R,Lane Brown R,Robinson P R.Light-dependent phosphorylation of the carboxy tail of mouse melanopsin.Cell Mol Life Sci,2012,69:1551-1562
35 Cameron E G,Robinson P R.β-arrestin-dependent deactivation of mouse melanopsin.PLoS One,2014,9:ell3138
36 Godement P,Salailn J,Imbert M.Prenatal and postnatal development of retinogeniculate and retinocollicular projections in the mouse.J Comp Neurol,1984,230:552-575
37 Johnson R F,Morin L P,Moore R Y.Retinohypothalamic projections in the hamster and rat demonstrated using cholera toxin.Brain Res,1988,462:301-312
38 Pickard G E.The afferent connections of the suprachiasmatic nucleus of the golden hamster with emphasis on the retinohypothalamic projection.J Comp Neurol,1982,211:65-83
39 Moore R Y,Speh J C,Patrick Card J.The retinohypothalamic tract originates from a distinct subset of retinal ganglion cells.J Comp Neurol,1995,352:351-366
40 Berson D M,Dunn F A,Takao M.Phototransduction by retinal ganglion cells that set the circadian clock.Science,2002,295:1070-1073
41 Hattar S,Kumar M,Park A,et al.Central projections of melanopsin-expressing retinal ganglion cells in the mouse.J Comp Neurol,2006,497:326-349
42 Baver S B,Pickard G E,Sollars P J,et al.Two types of melanopsin retinal ganglion cell differentially innervate the hypothalamic suprachiasmatic nucleus and the olivary pretectal nucleus.Eur J Neurosci,2008,27:1763-1770
43 Hannibal J,Hindersson P,Knudsen S M,et al.The photopigment melanopsin is exclusively present in pituitary adenylate cyclase-activating polypeptide-containing retinal ganglion cells of the retinohypothalamic tract.J Neurosci,2002,22:RC191
44 Berson D.Strange vision:ganglion cells as circadian photoreceptors.Trends Neurosci,2003,26:314-320
45 Hirota T,Fukada Y.Resetting mechanism of central and peripheral circadian clocks in mammals.Zoolog Sci,2004,21:359-368
46 MorinL P,Blanchard J H,Provencio I.Retinal ganglion cell projections to the hamster suprachiasmatic nucleus,intergeniculate leaflet,and visual midbrain:bifurcation and melanopsin immunoreactivity.J Comp Neurol,2003,465:401-416
47 Harrington M E.The ventral lateral geniculate nucleus and the intergeniculate leaflet:interrelated structures in the visual and circadian systems.Neurosci Biobehav Rev,1997,21:705-727
48 Trejo L J,Cicerone C M.Cells in the pretectal olivary nucleus are in the pathway forthe direct light reflex of the pupil in the rat.Brain Res,1984,300:49-62
49 Gamlin P D R,Zhang H,Clarke R J.Luminance neurons in the pretectal olivary nucleus mediate the pupillary light reflex in the rhesus monkey.Exp Brain Res,1995,106:169
50 Young M J,Lund R D.The anatomical substrates subserving the pupillary light reflex in rats:origin of the consensual pupillary response.Neuroscience,1994,62:481-496
51 Muscat L,Huberman A D,Jordan C L,et al.Crossed and uncrossed retinal projections to the hamster circadian system.J Comp Neurol,2003,466:513-524
52 Hannibal J,Fahrenkrug J.Target areas innervated by PACAP-immunoreactive retinal ganglion cells.Cell Tissue Res,2004,316:99-113
53 Ecker J L,Dumitrescu O N,Wong K Y,et al.Melanopsin-expressing retinal ganglion-cell photoreceptors:cellular diversity and role in pattern vision.Neuron,2010,67:49-60
54 Hannibal J,Kankipati L,Strang C E,et al.Central projections of intrinsically photosensitive retinal ganglion cells in the macaque monkey.J Comp Neurol,2014,522:2231-2248
55 Hannibal J,Hindersson P,Ostergaard J,et al.Melanopsin is expressed in pacap-containing retinal ganglion cells of the human retinohypothalamic tract.Invest Ophthalmol Vis Sci,2004,45:4202-4209
56 Lucas R J,Douglas R H,Foster R G.Characterization of an ocular photopigment capable of driving pupillary constriction in mice.Nat Neurosci,2001,4:621-626
57 Semo M,Peirson S,Lupi D,et al.Melanopsin retinal ganglion cells and the maintenance of circadian and pupillary responses to light in aged rodless/coneless(rd/rd cl)mice.Eur J Neurosci,2003,17:1793-1801
58 Reppert S M,Weaver D R.Coordination of circadian timing in mammals.Nature,2002,418:935-941
59 Panda S,Sato T K,Castrucci A M,et al.Melanopsin(Opn4)requirement for normal light-induced circadian phase shifting.Science,2002,298:2213-2216
60 Roecklein K A,Rohan K J,Duncan WC,et al.A missense variant(P10L)of the melanopsin(OPN4)gene in seasonal affective disorder.J Affect Disord,2009,114:279-285
61 Abrahamson E E,Moore R Y.Suprachiasmatic nucleus in the mouse:retinal innervation,intrinsic organization and efferent projections.Brain Res,2001,916:172-191
62 Luo A H,Aston-Jones G.Circuit projection from suprachiasmatic nucleus to ventral tegmental area:a novel circadian output pathway.Eur J Neurosci,2009,29:748-760
63 Altimus C M,G(u|¨)ler A D,Villa K L,et al.Rods-cones and melanopsin detect light and dark to modulate sleep independent of image formation.Proc Natl Acad Sci USA,2008,105:19998-20003
64 Lupi D,Oster H,Thompson S,et al.The acute light-induction of sleep is mediated by OPN4-based photoreception.Nat Neurosci,2008,11:1068-1073
65 Kalsbeek A,Palm I F,La Fleur S E,et al.SCN outputs and the hypothalamic balance of life.J Biol Rhythms,2006,21:458-469
66 Czeisler C A,Shanahan T L,Klerman E B,et al.Suppression of melatonin secretion in some blind patients by exposure to bright light.N Engl J Med,1995,332:6-11
67 Srinivasan V,Smits M,Spence W,et al.Melatonin in mood disorders.World J Biol Psychiatry,2006,7:138-151
68 LeGates T A,Fernandez D C,Hattar S.Light as a central modulator of circadian rhythms,sleep and affect.Nat Rev Neurosci,2014,15:443-454
69 Perlis M L,Giles D E,Buysse D J,et al.Self-reported sleep disturbance as a prodromal symptom in recurrent depression.J Affect Disord,1997,42:209-212
70 Brown R E,Basheer R,McKenna J T,et al.Control of sleep and wakefulness.Physiol Rev,2012,92:1087-1187
71 Perrin F,Peigneux P,Fuchs S,et al.Nonvisual responses to light exposure in the human brain during the circadian night.Curr Biol,2004,14:1842-1846
72 Vandewalle G,Maquet P,Dijk D J.Light as a modulator of cognitive brain function.Trends Cogn Sci,2009,13:429-438
73 Vandewalle G,Collignon O,Hull J T,et al.Blue light stimulates cognitive brain activity in visually blind individuals.J Cogn Neurosci,2013,25:2072-2085
74 Vandewalle G,Gais S,Schabus M,et al.Wavelength-dependent modulation of brain responses to a working memory task by daytime light exposure.Cerebral Cortex,2007,17:2788-2795
75 Bonmati-Carrion M A,Arguelles-Prieto R,Martinez-Madrid M J,et al.Protecting the melatonin rhythm through circadian healthy light exposure.Int J Mol Sci,2014,15:23448-23500
76 Haus E L,Smolensky M H.Shift work and cancer risk:potential mechanistic roles of circadian disruption,light at night,and sleep deprivation.Sleep Med Rev,2013,17:273-284
77 Stephenson R,Lim J,Famina S,et al.Sleep-wake behavior in the rat.J Biol Rhythms,2012,27:490-501
78 Lewy A J,Lefler B J,Emens J S,et al.The circadian basis of winter depression.Proc Natl Acad Sci USA,2006,103:7414-7419