KEGG   PATHWAY: epz04713
Entry
epz04713                    Pathway                                
Name
Circadian entrainment - Equus przewalskii (Przewalski's horse)
Description
Circadian entrainment is a fundamental property by which the period of the internal biological clock is entrained by recurring exogenous signals, such that the organism's endocrine and behavioral rhythms are synchronized to environmental cues. In mammals, a master clock is located in the suprachiasmatic nuclei (SCN) of the hypothalamus and may synchronize circadian oscillators in peripheral tissues. Light signal is the dominant synchronizer for master SCN clock. Downstream from the retina, glutamate and PACAP are released and trigger the activation of signal transduction cascades, including CamKII and nNOS activity, cAMP- and cGMP-dependent protein kinases, and mitogen-activated protein kinase (MAPK). Of non-photic entrainment, important phase shifting capabilities have been found for melatonin, which inhibits light-induced phase shifts through inhibition of adenylate cyclase (AC). Multiple entrainment pathways converge into CREB regulation. In turn, phosphorylated CREB activates clock gene expression.
Class
Organismal Systems; Environmental adaptation
Pathway map
epz04713  Circadian entrainment
epz04713

Other DBs
GO: 0009649
Organism
Equus przewalskii (Przewalski's horse) [GN:epz]
Gene
103563588  GRIN1; glutamate ionotropic receptor NMDA type subunit 1 [KO:K05208]
103555146  GRIN2A; glutamate ionotropic receptor NMDA type subunit 2A [KO:K05209]
103549904  GRIN2B; glutamate ionotropic receptor NMDA type subunit 2B [KO:K05210]
103546604  GRIN2C; glutamate ionotropic receptor NMDA type subunit 2C [KO:K05211]
103541380  GRIN2D; glutamate ionotropic receptor NMDA type subunit 2D [KO:K05212]
103548885  CALML5; calmodulin like 5 [KO:K02183]
103548886  CALML3; calmodulin like 3 [KO:K02183]
103549905  CALM1; calmodulin 1 [KO:K02183]
103558105  CALM2; calmodulin 2 [KO:K02183]
103564053  calmodulin-like protein 4 [KO:K02183]
103544963  calmodulin-alpha-like [KO:K02183]
103567415  CALML6; calmodulin like 6 [KO:K02183]
103549465  CAMK2G; calcium/calmodulin dependent protein kinase II gamma [KO:K04515] [EC:2.7.11.17]
103549616  CAMK2D; calcium/calmodulin dependent protein kinase II delta [KO:K04515] [EC:2.7.11.17]
103549121  CAMK2A; calcium/calmodulin dependent protein kinase II alpha [KO:K04515] [EC:2.7.11.17]
103554952  CAMK2B; calcium/calmodulin dependent protein kinase II beta [KO:K04515] [EC:2.7.11.17]
103547950  NOS1; nitric oxide synthase 1 [KO:K13240] [EC:1.14.13.39]
103547261  MAPK3; mitogen-activated protein kinase 3 [KO:K04371] [EC:2.7.11.24]
103562408  MAPK1; mitogen-activated protein kinase 1 [KO:K04371] [EC:2.7.11.24]
103558014  RPS6KA5; ribosomal protein S6 kinase A5 [KO:K04445] [EC:2.7.11.1]
103548908  CREB1; cAMP responsive element binding protein 1 [KO:K05870]
103550297  ITPR1; inositol 1,4,5-trisphosphate receptor type 1 [KO:K04958]
103564724  ITPR3; inositol 1,4,5-trisphosphate receptor type 3 [KO:K04960]
103541052  RYR1; ryanodine receptor 1 [KO:K04961]
103554178  RYR2; ryanodine receptor 2 [KO:K04962]
103548546  RYR3; ryanodine receptor 3 [KO:K04963]
103561275  carboxyl-terminal PDZ ligand of neuronal nitric oxide synthase protein-like [KO:K16513]
103552141  guanine nucleotide-binding protein G(i) subunit alpha-1-like [KO:K04630]
103553052  GNAI3; G protein subunit alpha i3 [KO:K04630]
103557343  GNAI2; G protein subunit alpha i2 [KO:K04630]
103555391  GNAO1; G protein subunit alpha o1 [KO:K04534]
103567413  GNB1; G protein subunit beta 1 [KO:K04536]
103555641  GNB2; G protein subunit beta 2 [KO:K04537]
103547020  GNB3; G protein subunit beta 3 [KO:K07825]
103551737  GNB4; G protein subunit beta 4 [KO:K04538]
103548465  GNB5; G protein subunit beta 5 [KO:K04539]
103543306  guanine nucleotide-binding protein G(I)/G(S)/G(O) subunit gamma-2 [KO:K07826]
103558941  GNG3; G protein subunit gamma 3 [KO:K04540]
103547680  GNG4; G protein subunit gamma 4 [KO:K04541]
103557283  GNG5; G protein subunit gamma 5 [KO:K04542]
103542500  GNG8; G protein subunit gamma 8 [KO:K04544]
103542169  GNG11; G protein subunit gamma 11 [KO:K04546]
103564847  GNG12; G protein subunit gamma 12 [KO:K04347]
103542772  guanine nucleotide-binding protein G(I)/G(S)/G(O) subunit gamma-7 [KO:K04347]
103565302  GNG13; G protein subunit gamma 13 [KO:K04547]
103568001  GNGT1; G protein subunit gamma transducin 1 [KO:K04548]
103551067  GNGT2; G protein subunit gamma transducin 2 [KO:K04549]
103549680  GRIA1; glutamate ionotropic receptor AMPA type subunit 1 [KO:K05197]
103547313  GRIA2; glutamate ionotropic receptor AMPA type subunit 2 [KO:K05198]
103557390  GRIA3; glutamate ionotropic receptor AMPA type subunit 3 [KO:K05199]
103540802  GRIA4; glutamate ionotropic receptor AMPA type subunit 4 [KO:K05200]
103549566  CACNA1G; calcium voltage-gated channel subunit alpha1 G [KO:K04854]
103565287  CACNA1I; calcium voltage-gated channel subunit alpha1 I [KO:K04856]
103563075  ADCYAP1; adenylate cyclase activating polypeptide 1 [KO:K05262]
103559362  ADCYAP1R1; ADCYAP receptor type I [KO:K04587]
103565402  guanine nucleotide-binding protein G(s) subunit alpha [KO:K04632]
103547036  neuroendocrine secretory protein 55 [KO:K04632]
103550430  ADCY1; adenylate cyclase 1 [KO:K08041] [EC:4.6.1.1]
103562505  adenylate cyclase type 2-like [KO:K08042] [EC:4.6.1.1]
103547823  ADCY3; adenylate cyclase 3 [KO:K08043] [EC:4.6.1.1]
103561050  ADCY4; adenylate cyclase 4 [KO:K08044] [EC:4.6.1.1]
103563830  ADCY5; adenylate cyclase 5 [KO:K08045] [EC:4.6.1.1]
103563634  ADCY6; adenylate cyclase 6 [KO:K08046] [EC:4.6.1.1]
103556685  ADCY7; adenylate cyclase 7 [KO:K08047] [EC:4.6.1.1]
103565019  adenylate cyclase type 8-like [KO:K08048] [EC:4.6.1.1]
103567623  ADCY8; adenylate cyclase 8 [KO:K08048] [EC:4.6.1.1]
103553014  ADCY9; adenylate cyclase 9 [KO:K08049] [EC:4.6.1.1]
103551549  ADCY10; adenylate cyclase 10, soluble [KO:K11265] [EC:4.6.1.1]
103555495  adenylate cyclase type 10-like [KO:K11265] [EC:4.6.1.1]
103557289  PRKACB; protein kinase cAMP-activated catalytic subunit beta [KO:K04345] [EC:2.7.11.11]
103565953  PRKACA; protein kinase cAMP-activated catalytic subunit alpha [KO:K04345] [EC:2.7.11.11]
103563826  guanylate cyclase soluble subunit alpha-2 [KO:K12318] [EC:4.6.1.2]
103562610  GUCY1A3; guanylate cyclase 1 soluble subunit alpha [KO:K12318] [EC:4.6.1.2]
103551994  guanylate cyclase soluble subunit beta-2-like [KO:K12319] [EC:4.6.1.2]
103562608  GUCY1B3; guanylate cyclase 1 soluble subunit beta [KO:K12319] [EC:4.6.1.2]
103549039  PRKG2; protein kinase, cGMP-dependent, type II [KO:K19477] [EC:2.7.11.12]
103560754  voltage-dependent L-type calcium channel subunit alpha-1D [KO:K04851]
103548037  MTNR1B; melatonin receptor 1B [KO:K04286]
103549958  MTNR1A; melatonin receptor 1A [KO:K04285]
103553450  GNAQ; G protein subunit alpha q [KO:K04634]
103562440  PLCB1; phospholipase C beta 1 [KO:K05858] [EC:3.1.4.11]
103559733  PLCB4; phospholipase C beta 4 [KO:K05858] [EC:3.1.4.11]
103561633  PLCB2; phospholipase C beta 2 [KO:K05858] [EC:3.1.4.11]
103542193  PLCB3; phospholipase C beta 3 [KO:K05858] [EC:3.1.4.11]
103558165  PRKCB; protein kinase C beta [KO:K19662] [EC:2.7.11.13]
103556196  PRKCG; protein kinase C gamma [KO:K19663] [EC:2.7.11.13]
103555321  KCNJ3; potassium voltage-gated channel subfamily J member 3 [KO:K04997]
103551404  KCNJ5; potassium voltage-gated channel subfamily J member 5 [KO:K04999]
103564035  KCNJ6; potassium voltage-gated channel subfamily J member 6 [KO:K05000]
103551152  KCNJ9; potassium voltage-gated channel subfamily J member 9 [KO:K05002]
103564278  PER1; period circadian clock 1 [KO:K21944]
103559632  PER2; period circadian clock 2 [KO:K02633]
103549573  PER3; period circadian clock 3 [KO:K21945]
103548618  FOS; Fos proto-oncogene, AP-1 transcription factor subunit [KO:K04379]
Compound
C00025  L-Glutamate
C00076  Calcium cation
C00165  Diacylglycerol
C00238  Potassium cation
C00533  Nitric oxide
C00575  3',5'-Cyclic AMP
C00942  3',5'-Cyclic GMP
C01330  Sodium cation
C01598  Melatonin
Reference
  Authors
Brown TM, Piggins HD
  Title
Electrophysiology of the suprachiasmatic circadian clock.
  Journal
Prog Neurobiol 82:229-55 (2007)
DOI:10.1016/j.pneurobio.2007.05.002
Reference
  Authors
Dickson L, Finlayson K
  Title
VPAC and PAC receptors: From ligands to function.
  Journal
Pharmacol Ther 121:294-316 (2009)
DOI:10.1016/j.pharmthera.2008.11.006
Reference
  Authors
Golombek DA, Rosenstein RE
  Title
Physiology of circadian entrainment.
  Journal
Physiol Rev 90:1063-102 (2010)
DOI:10.1152/physrev.00009.2009
Reference
  Authors
Dubocovich ML, Markowska M
  Title
Functional MT1 and MT2 melatonin receptors in mammals.
  Journal
Endocrine 27:101-10 (2005)
DOI:10.1385/ENDO:27:2:101
Reference
  Authors
Dubocovich ML, Delagrange P, Krause DN, Sugden D, Cardinali DP, Olcese J
  Title
International Union of Basic and Clinical Pharmacology. LXXV. Nomenclature, classification, and pharmacology of G protein-coupled melatonin receptors.
  Journal
Pharmacol Rev 62:343-80 (2010)
DOI:10.1124/pr.110.002832
Reference
  Authors
Hardeland R
  Title
Melatonin: signaling mechanisms of a pleiotropic agent.
  Journal
Biofactors 35:183-92 (2009)
DOI:10.1002/biof.23
Reference
  Authors
Golombek DA, Ferreyra GA, Agostino PV, Murad AD, Rubio MF, Pizzio GA, Katz ME, Marpegan L, Bekinschtein TA
  Title
From light to genes: moving the hands of the circadian clock.
  Journal
Front Biosci 8:s285-93 (2003)
DOI:10.2741/1038
Reference
  Authors
Mertens I, Husson SJ, Janssen T, Lindemans M, Schoofs L
  Title
PACAP and PDF signaling in the regulation of mammalian and insect circadian rhythms.
  Journal
Peptides 28:1775-83 (2007)
DOI:10.1016/j.peptides.2007.05.005
Reference
  Authors
Kim DY, Choi HJ, Kim JS, Kim YS, Jeong DU, Shin HC, Kim MJ, Han HC, Hong SK, Kim YI
  Title
Voltage-gated calcium channels play crucial roles in the glutamate-induced phase shifts of the rat suprachiasmatic circadian clock.
  Journal
Eur J Neurosci 21:1215-22 (2005)
DOI:10.1111/j.1460-9568.2005.03950.x
Reference
  Authors
Agostino PV, Ferreyra GA, Murad AD, Watanabe Y, Golombek DA
  Title
Diurnal, circadian and photic regulation of calcium/calmodulin-dependent kinase II and neuronal nitric oxide synthase in the hamster suprachiasmatic nuclei.
  Journal
Neurochem Int 44:617-25 (2004)
DOI:10.1016/j.neuint.2003.09.005
Reference
  Authors
Ferreyra GA, Golombek DA
  Title
Rhythmicity of the cGMP-related signal transduction pathway in the mammalian circadian system.
  Journal
Am J Physiol Regul Integr Comp Physiol 280:R1348-55 (2001)
DOI:10.1152/ajpregu.2001.280.5.R1348
Reference
  Authors
Butcher GQ, Lee B, Cheng HY, Obrietan K
  Title
Light stimulates MSK1 activation in the suprachiasmatic nucleus via a PACAP-ERK/MAP kinase-dependent mechanism.
  Journal
J Neurosci 25:5305-13 (2005)
DOI:10.1523/JNEUROSCI.4361-04.2005
Reference
  Authors
Hamada T, Niki T, Ziging P, Sugiyama T, Watanabe S, Mikoshiba K, Ishida N
  Title
Differential expression patterns of inositol trisphosphate receptor types 1 and 3 in the rat suprachiasmatic nucleus.
  Journal
Brain Res 838:131-5 (1999)
DOI:10.1016/S0006-8993(99)01719-9
Reference
  Authors
Diaz-Munoz M, Dent MA, Granados-Fuentes D, Hall AC, Hernandez-Cruz A, Harrington ME, Aguilar-Roblero R
  Title
Circadian modulation of the ryanodine receptor type 2 in the SCN of rodents.
  Journal
Neuroreport 10:481-6 (1999)
DOI:10.1097/00001756-199902250-00007
Reference
  Authors
Fang M, Jaffrey SR, Sawa A, Ye K, Luo X, Snyder SH
  Title
Dexras1: a G protein specifically coupled to neuronal nitric oxide synthase via CAPON.
  Journal
Neuron 28:183-93 (2000)
DOI:10.1016/S0896-6273(00)00095-7
Reference
  Authors
Takahashi H, Umeda N, Tsutsumi Y, Fukumura R, Ohkaze H, Sujino M, van der Horst G, Yasui A, Inouye ST, Fujimori A, Ohhata T, Araki R, Abe M
  Title
Mouse dexamethasone-induced RAS protein 1 gene is expressed in a circadian rhythmic manner in the suprachiasmatic nucleus.
  Journal
Brain Res Mol Brain Res 110:1-6 (2003)
DOI:10.1016/S0169-328X(02)00543-0
Reference
  Authors
Cheng HY, Dziema H, Papp J, Mathur DP, Koletar M, Ralph MR, Penninger JM, Obrietan K
  Title
The molecular gatekeeper Dexras1 sculpts the photic responsiveness of the mammalian circadian clock.
  Journal
J Neurosci 26:12984-95 (2006)
DOI:10.1523/JNEUROSCI.4253-06.2006
Related
pathway
epz04010  MAPK signaling pathway
epz04710  Circadian rhythm
KO pathway
ko04713   
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