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Walton, Z., Patel, C.H., Brooks, R.C., Yu, Y., Ibrahim-Hashim, A., Riddle, M., Porcu, A., Tianying, J., Ecker, B.L., Tameire, F., Koumenis, C., Weeraratna, A.T., Welsh, D.K., Gillies, R., Alwine, J.C., Zhang, L., Powell, J.D., and Dang, C.. Acid suspends the circadian clock in hypoxia through inhibition of mTOR. Cell, 28 June 2018.

Photoperiod-induced neuroplasticity in the circadian system.
Porcu, A, Riddle, M, Dulcis, D, Welsh, DK. 
Neural Plasticity, 2018 Feb 28. Review. 

Circadian Clocks as Modulators of Metabolic Comorbidity in Psychiatric Disorders.
Barandas R, Landgraf D, McCarthy MJ, Welsh DK.
Curr Psychiatry Rep. 2015 Dec;17(12):98. doi: 10.1007/s11920-015-0637-2. Review.

Photoperiodic and circadian bifurcation theories of depression and mania.
Kripke DF, Elliott JA, Welsh DK, Youngstedt SD.
F1000Res. 2015 May 6;4:107. doi: 10.12688/f1000research.6444.1. eCollection 2015. Review.

Circadian clock and stress interactions in the molecular biology of psychiatric disorders.
Landgraf D, McCarthy MJ, Welsh DK.
Curr Psychiatry Rep. 2014 Oct;16(10):483. doi: 10.1007/s11920-014-0483-7. Review.

The role of the circadian clock in animal models of mood disorders.
Landgraf D, McCarthy MJ, Welsh DK.
Behav Neurosci. 2014 Jun;128(3):344-59. doi: 10.1037/a0036029. Epub 2014 Mar 24. Review.

Cellular circadian clocks in mood disorders.
McCarthy MJ, Welsh DK.
J Biol Rhythms. 2012 Oct;27(5):339-52. Review.

Suprachiasmatic nucleus: cell autonomy and network properties.
Welsh DK, Takahashi JS, Kay SA.
Annu Rev Physiol. 2010;72:551-77. doi: 10.1146/annurev-physiol-021909-135919. Review.

Exploring spatiotemporal organization of SCN circuits.
Yan L, Karatsoreos I, Lesauter J, Welsh DK, Kay S, Foley D, Silver R.
Cold Spring Harb Symp Quant Biol. 2007;72:527-41. doi: 10.1101/sqb.2007.72.037. Review.

Beyond intuitive modeling: combining biophysical models with innovative experiments to move the circadian clock field forward.
Forger D, Gonze D, Virshup D, Welsh DK.
J Biol Rhythms. 2007 Jun;22(3):200-10. Review.

Bioluminescence imaging in living organisms.
Welsh DK, Kay SA.
Curr Opin Biotechnol. 2005 Feb;16(1):73-8. Review.

The role of chronobiology in sleep disorders medicine.
Monk TH, Welsh DK.
Sleep Med Rev. 2003 Dec;7(6):455-73. Review.

Original Research


Cellular circadian oscillators in the suprachiasmatic nucleus remain coupled in the absence of connexin-36.
Diemer T, Landgraf D, Noguchi T, Pan H, Moreno JL, Welsh DK.
Neuroscience. 2017 Aug 15;357:1-11. doi: 10.1016/j.neuroscience.2017.05.037. Epub 2017 May 31.

Enhancing circadian clock function in cancer cells inhibits tumor growth.
Kiessling S, Beaulieu-Laroche L, Blum ID, Landgraf D, Welsh DK, Storch KF, Labrecque N, Cermakian N.
BMC Biol. 2017 Feb 14;15(1):13. doi: 10.1186/s12915-017-0349-7.

Circadian alterations during early stages of Alzheimer's disease are associated with aberrant cycles of DNA methylation in BMAL1.
Cronin P, McCarthy MJ, Lim ASP, Salmon DP, Galasko D, Masliah E, De Jager PL, Bennett DA, Desplats P.
Alzheimers Dement. 2017 Jun;13(6):689-700. doi: 10.1016/j.jalz.2016.10.003. Epub 2016 Nov 22.

Cardiomyocyte Circadian Oscillations Are Cell-Autonomous, Amplified by β-Adrenergic Signaling, and Synchronized in Cardiac Ventricle Tissue.
Beesley S, Noguchi T, Welsh DK.
PLoS One. 2016 Jul 26;11(7):e0159618. doi: 10.1371/journal.pone.0159618. eCollection 2016.

Functional Contributions of Strong and Weak Cellular Oscillators to Synchrony and Light-shifted Phase Dynamics.
Roberts L, Leise TL, Welsh DK, Holmes TC.
J Biol Rhythms. 2016 Aug;31(4):337-51. doi: 10.1177/0748730416649550. Epub 2016 May 24.

Disinhibition of the extracellular-signal-regulated kinase restores the amplification of circadian rhythms by lithium in cells from bipolar disorder patients.
McCarthy MJ, Wei H, Landgraf D, Le Roux MJ, Welsh DK.
Eur Neuropsychopharmacol. 2016 Aug;26(8):1310-9. doi: 10.1016/j.euroneuro.2016.05.003. Epub 2016 May 20.

The Pharmacogenomics of Bipolar Disorder study (PGBD): identification of genes for lithium response in a prospective sample.
Oedegaard KJ, Alda M, Anand A, Andreassen OA, Balaraman Y, Berrettini WH, Bhattacharjee A, Brennand KJ, Burdick KE, Calabrese JR, Calkin CV, Claasen A, Coryell WH, Craig D, DeModena A, Frye M, Gage FH, Gao K, Garnham J, Gershon E, Jakobsen P, Leckband SG, McCarthy MJ, McInnis MG, Maihofer AX, Mertens J, Morken G, Nievergelt CM, Nurnberger J, Pham S, Schoeyen H, Shekhtman T, Shilling PD, Szelinger S, Tarwater B, Yao J, Zandi PP, Kelsoe JR.
BMC Psychiatry. 2016 May 5;16:129. doi: 10.1186/s12888-016-0732-x.

Genetic Disruption of Circadian Rhythms in the Suprachiasmatic Nucleus Causes Helplessness, Behavioral Despair, and Anxiety-like Behavior in Mice.
Landgraf D, Long JE, Proulx CD, Barandas R, Malinow R, Welsh DK.
Biol Psychiatry. 2016 Dec 1;80(11):827-835. doi: 10.1016/j.biopsych.2016.03.1050. Epub 2016 Mar 10.

The mood stabilizer valproic acid opposes the effects of dopamine on circadian rhythms.
Landgraf D, Joiner WJ, McCarthy MJ, Kiessling S, Barandas R, Young JW, Cermakian N, Welsh DK.
Neuropharmacology. 2016 Aug;107:262-70. doi: 10.1016/j.neuropharm.2016.03.047. Epub 2016 Mar 28.

Lithium effects on circadian rhythms in fibroblasts and suprachiasmatic nucleus slices from Cry knockout mice.
Noguchi T, Lo K, Diemer T, Welsh DK.
Neurosci Lett. 2016 Apr 21;619:49-53. doi: 10.1016/j.neulet.2016.02.030. Epub 2016 Feb 27.

NPAS2 Compensates for Loss of CLOCK in Peripheral Circadian Oscillators.
Landgraf D, Wang LL, Diemer T, Welsh DK.
PLoS Genet. 2016 Feb 19;12(2):e1005882. doi: 10.1371/journal.pgen.1005882. eCollection 2016 Feb.


Calcium channel genes associated with bipolar disorder modulate lithium's amplification of circadian rhythms.
McCarthy MJ, Le Roux MJ, Wei H, Beesley S, Kelsoe JR, Welsh DK.
Neuropharmacology. 2016 Feb;101:439-48. doi: 10.1016/j.neuropharm.2015.10.017. Epub 2015 Oct 22.

Depression-like behaviour in mice is associated with disrupted circadian rhythms in nucleus accumbens and periaqueductal grey.
Landgraf D, Long JE, Welsh DK.
Eur J Neurosci. 2016 May;43(10):1309-20. doi: 10.1111/ejn.13085. Epub 2015 Nov 6.

Dissociation of learned helplessness and fear conditioning in mice: a mouse model of depression.
Landgraf D, Long J, Der-Avakian A, Streets M, Welsh DK.
PLoS One. 2015 Apr 30;10(4):e0125892. doi: 10.1371/journal.pone.0125892. eCollection 2015.

Light evokes rapid circadian network oscillator desynchrony followed by gradual phase retuning of synchrony.
Roberts L, Leise TL, Noguchi T, Galschiodt AM, Houl JH, Welsh DK, Holmes TC.
Curr Biol. 2015 Mar 30;25(7):858-67. doi: 10.1016/j.cub.2015.01.056. Epub 2015 Mar 5.

Towards the clinical implementation of pharmacogenetics in bipolar disorder.
Salloum NC, McCarthy MJ, Leckband SG, Kelsoe JR.
BMC Med. 2014 May 30;12:90. doi: 10.1186/1741-7015-12-90.

Cell type-specific functions of period genes revealed by novel adipocyte and hepatocyte circadian clock models.
Ramanathan C, Xu H, Khan SK, Shen Y, Gitis PJ, Welsh DK, Hogenesch JB, Liu AC.
PLoS Genet. 2014 Apr 3;10(4):e1004244. doi: 10.1371/journal.pgen.1004244. eCollection 2014 Apr.

Oxidative stress: a link between cardiovascular disease and psychiatric illness?
McCarthy MJ.
Acta Psychiatr Scand. 2014 Sep;130(3):161-2. doi: 10.1111/acps.12269. Epub 2014 Mar 24.

Genetic and clinical factors predict lithium's effects on PER2 gene expression rhythms in cells from bipolar disorder patients.
McCarthy MJ, Wei H, Marnoy Z, Darvish RM, McPhie DL, Cohen BM, Welsh DK.
Transl Psychiatry. 2013 Oct 22;3:e318. doi: 10.1038/tp.2013.90.

Fibroblast PER2 circadian rhythmicity depends on cell density.
Noguchi T, Wang LL, Welsh DK.
J Biol Rhythms. 2013 Jun;28(3):183-92. doi: 10.1177/0748730413487494.

Circadian clock period inversely correlates with illness severity in cells from patients with alcohol use disorders.
McCarthy MJ, Fernandes M, Kranzler HR, Covault JM, Welsh DK.
Alcohol Clin Exp Res. 2013 Aug;37(8):1304-10. doi: 10.1111/acer.12106. Epub 2013 Mar 29.

Cry1-/- circadian rhythmicity depends on SCN intercellular coupling.
Evans JA, Pan H, Liu AC, Welsh DK.
J Biol Rhythms. 2012 Dec;27(6):443-52. doi: 10.1177/0748730412461246.

Bayesian statistical analysis of circadian oscillations in fibroblasts.
Cohen AL, Leise TL, Welsh DK.
J Theor Biol. 2012 Dec 7;314:182-91. doi: 10.1016/j.jtbi.2012.08.038. Epub 2012 Sep 8.

Cellular bioluminescence imaging.
Welsh DK, Noguchi T.
Cold Spring Harb Protoc. 2012 Aug 1;2012(8). pii: pdb.top070607. doi: 10.1101/pdb.top070607. Erratum in: Cold Spring Harb Protoc. 2012 Sep;2012(9):1028. Cold Spring Harb Protoc. 2016;2016(3):pdb.err091751.

Identification of small molecule activators of cryptochrome.
Hirota T, Lee JW, St John PC, Sawa M, Iwaisako K, Noguchi T, Pongsawakul PY, Sonntag T, Welsh DK, Brenner DA, Doyle FJ 3rd, Schultz PG, Kay SA.
Science. 2012 Aug 31;337(6098):1094-7. doi: 10.1126/science.1223710. Epub 2012 Jul 12.

Fibroblast circadian rhythms of PER2 expression depend on membrane potential and intracellular calcium.
Noguchi T, Wang CW, Pan H, Welsh DK.
Chronobiol Int. 2012 Jul;29(6):653-64. doi: 10.3109/07420528.2012.679330.

Persistent cell-autonomous circadian oscillations in fibroblasts revealed by six-week single-cell imaging of PER2::LUC bioluminescence.
Leise TL, Wang CW, Gitis PJ, Welsh DK.
PLoS One. 2012;7(3):e33334. doi: 10.1371/journal.pone.0033334. Epub 2012 Mar 29.

A survey of genomic studies supports association of circadian clock genes with bipolar disorder spectrum illnesses and lithium response.
McCarthy MJ, Nievergelt CM, Kelsoe JR, Welsh DK.
PLoS One. 2012;7(2):e32091. doi: 10.1371/journal.pone.0032091. Epub 2012 Feb 22.

Functional genetic variation in the Rev-Erbα pathway and lithium response in the treatment of bipolar disorder.
McCarthy MJ, Nievergelt CM, Shekhtman T, Kripke DF, Welsh DK, Kelsoe JR.
Genes Brain Behav. 2011 Nov;10(8):852-61. doi: 10.1111/j.1601-183X.2011.00725.x. Epub 2011 Sep 2.

Polymorphisms in melatonin synthesis pathways: possible influences on depression.
Kripke DF, Nievergelt CM, Tranah GJ, Murray SS, McCarthy MJ, Rex KM, Parimi N, Kelsoe JR.
J Circadian Rhythms. 2011 Aug 9;9:8. doi: 10.1186/1740-3391-9-8.

Wavelet measurement suggests cause of period instability in mammalian circadian neurons.
Meeker K, Harang R, Webb AB, Welsh DK, Doyle FJ 3rd, Bonnet G, Herzog ED, Petzold LR.
J Biol Rhythms. 2011 Aug;26(4):353-62. doi: 10.1177/0748730411409863.

Characterization of orderly spatiotemporal patterns of clock gene activation in mammalian suprachiasmatic nucleus.
Foley NC, Tong TY, Foley D, Lesauter J, Welsh DK, Silver R.
Eur J Neurosci. 2011 May;33(10):1851-65. doi: 10.1111/j.1460-9568.2011.07682.x. Epub 2011 Apr 14.

Duplications of the neuropeptide receptor gene VIPR2 confer significant risk for schizophrenia.
Vacic V, McCarthy S, Malhotra D, Murray F, Chou HH, Peoples A, Makarov V, Yoon S, Bhandari A, Corominas R, Iakoucheva LM, Krastoshevsky O, Krause V, Larach-Walters V, Welsh DK, Craig D, Kelsoe JR, Gershon ES, Leal SM, Dell Aquila M, Morris DW, Gill M, Corvin A, Insel PA, McClellan J, King MC, Karayiorgou M, Levy DL, DeLisi LE, Sebat J.
Nature. 2011 Mar 24;471(7339):499-503. doi: 10.1038/nature09884. Epub 2011 Feb 23. Erratum in: Nature. 2011 Jun 2;474(7349):114.

Emergence of noise-induced oscillations in the central circadian pacemaker.
Ko CH, Yamada YR, Welsh DK, Buhr ED, Liu AC, Zhang EE, Ralph MR, Kay SA, Forger DB, Takahashi JS.
PLoS Biol. 2010 Oct 12;8(10):e1000513. doi: 10.1371/journal.pbio.1000513. Erratum in: PLoS Biol. 2010;8(10) doi: 10.1371/annotation/45e0bb8b-69bd-4ba0-bb9e-6641bce10fc6.

Internet monitoring of suicide risk in the population.
McCarthy MJ.
J Affect Disord. 2010 May;122(3):277-9. doi: 10.1016/j.jad.2009.08.015. Epub 2009 Sep 12.


Quantification of circadian rhythms in single cells.
Westermark PO, Welsh DK, Okamura H, Herzel H.
PLoS Comput Biol. 2009 Nov;5(11):e1000580. doi: 10.1371/journal.pcbi.1000580. Epub 2009 Nov 26.

Expression of the circadian clock gene Period2 in the hippocampus: possible implications for synaptic plasticity and learned behaviour.
Wang LM, Dragich JM, Kudo T, Odom IH, Welsh DK, O'Dell TJ, Colwell CS.
ASN Neuro. 2009 Jun 10;1(3). pii: e00012. doi: 10.1042/AN20090020.

A model of the cell-autonomous mammalian circadian clock.
Mirsky HP, Liu AC, Welsh DK, Kay SA, Doyle FJ 3rd.
Proc Natl Acad Sci U S A. 2009 Jul 7;106(27):11107-12. doi: 10.1073/pnas.0904837106. Epub 2009 Jun 19.

Redundant function of REV-ERBalpha and beta and non-essential role for Bmal1 cycling in transcriptional regulation of intracellular circadian rhythms.
Liu AC, Tran HG, Zhang EE, Priest AA, Welsh DK, Kay SA.
PLoS Genet. 2008 Feb 29;4(2):e1000023. doi: 10.1371/journal.pgen.1000023.

Reciprocity between phase shifts and amplitude changes in the mammalian circadian clock.
Pulivarthy SR, Tanaka N, Welsh DK, De Haro L, Verma IM, Panda S.
Proc Natl Acad Sci U S A. 2007 Dec 18;104(51):20356-61. Epub 2007 Dec 7.

Intercellular coupling confers robustness against mutations in the SCN circadian clock network.
Liu AC, Welsh DK, Ko CH, Tran HG, Zhang EE, Priest AA, Buhr ED, Singer O, Meeker K, Verma IM, Doyle FJ 3rd, Takahashi JS, Kay SA.
Cell. 2007 May 4;129(3):605-16.

VIP activates and couples clock cells. Focus on "Disrupted neuronal activity rhythms in the suprachiasmatic nucleus of vasoactive intestinal polypeptide-deficient mice".
Welsh DK.
J Neurophysiol. 2007 Mar;97(3):1885-6. Epub 2007 Jan 24.

Gate cells see the light.
Welsh DK.
J Biol Rhythms. 2007 Feb;22(1):26-8.

Feedback repression is required for mammalian circadian clock function.
Sato TK, Yamada RG, Ukai H, Baggs JE, Miraglia LJ, Kobayashi TJ, Welsh DK, Kay SA, Ueda HR, Hogenesch JB.
Nat Genet. 2006 Mar;38(3):312-9. Epub 2006 Feb 12.

Real-time reporting of circadian-regulated gene expression by luciferase imaging in plants and mammalian cells.
Welsh DK, Imaizumi T, Kay SA.
Methods Enzymol. 2005;393:269-88.


Bioluminescence imaging of individual fibroblasts reveals persistent, independently phased circadian rhythms of clock gene expression.
Welsh DK, Yoo SH, Liu AC, Takahashi JS, Kay SA.
Curr Biol. 2004 Dec 29;14(24):2289-95.

A sleep diary and questionnaire study of naturally short sleepers.
Monk TH, Buysse DJ, Welsh DK, Kennedy KS, Rose LR.
J Sleep Res. 2001 Sep;10(3):173-9.

Prior to 2000 

Gap junctions couple astrocytes but not neurons in dissociated cultures of rat suprachiasmatic nucleus.
Welsh DK, Reppert SM.
Brain Res. 1996 Jan 8;706(1):30-6.

Individual neurons dissociated from rat suprachiasmatic nucleus express independently phased circadian firing rhythms.
Welsh DK, Logothetis DE, Meister M, Reppert SM.
Neuron. 1995 Apr;14(4):697-706.

Lithium lengthens circadian period in a diurnal primate, Saimiri sciureus.
Welsh DK, Moore-Ede MC.
Biol Psychiatry. 1990 Jul 15;28(2):117-26.

Aluminum fluoride reveals a phosphoinositide system within the suprachiasmatic region of rat hypothalamus.
Nadakavukaren JJ, Welsh DK, Reppert SM.
Brain Res. 1990 Jan 22;507(2):181-8.

Effect of age on the circadian pattern of sleep and wakefulness in the mouse.
Welsh DK, Richardson GS, Dement WC.
J Gerontol. 1986 Sep;41(5):579-86.

Precision of circadian wake and activity onset timing in the mouse.
Welsh DK, Engel EM, Richardson GS, Dement WC.
J Comp Physiol A. 1986 Jun;158(6):827-34. Erratum in: J Comp Physiol [A] 1986 Oct;159(4):577. Engle EM [corrected to Engel EM].

Regular 48-hour cycling of sleep duration and mood in a 35-year-old woman: use of lithium in time isolation.
Welsh DK, Nino-Murcia G, Gander PH, Keenan S, Dement WC.
Biol Psychiatry. 1986 May;21(5-6):527-37.

A circadian rhythm of hippocampal theta activity in the mouse.
Welsh DK, Richardson GS, Dement WC.
Physiol Behav. 1985 Oct;35(4):533-8.