David Welsh

Title(s)Professor, Psychiatry
SchoolHealth Sciences
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    Dr. Welsh earned his undergraduate degree at Stanford University, where he began a long-standing interest in circadian rhythms and sleep. In graduate work at Harvard Medical School, he discovered that individual brain cells can generate circadian rhythms. He received a Ph.D. in Neurobiology and an M.D. from Harvard in 1997, then completed a residency in general psychiatry at Western Psychiatric Institute and Clinic, in Pittsburgh. After two years as a postdoctoral research fellow, he joined the faculty of the UCSD Department of Psychiatry in 2003. He is currently an Associate Professor in the Department, and a Staff Psychiatrist at the VA San Diego Healthcare System.

    Research Interests

    In mammals, a hierarchical system of circadian clock cells in the brain and throughout the body orchestrates daily patterns of physiology and behavior. These daily patterns persist under constant conditions as "circadian rhythms". Dr. Welsh studies circadian rhythms in single cells using bioluminescence imaging to monitor clock gene expression. He is interested in the autonomy, heterogeneity, and coupling of cellular circadian clocks, particularly the "master" clock cells of the brain, the neurons of the suprachiasmatic nucleus. He is also interested in how defects in these mechanisms may contribute to sleep and circadian rhythm disorders in humans.

    Clinical Focus

    Dr. Welsh is a Staff Psychiatrist at the VA San Diego Healthcare System. He practices general adult psychiatry in the mood disorders clinic.

    Collapse Research 
    Collapse Research Activities and Funding
    Circadian dysfunction and Epigenetic alterations in the pathogenesis of Alzheimer's Disease
    NIH/NIA R01AG061831Feb 1, 2019 - Nov 30, 2023
    Role: Co-Principal Investigator
    Cellular Circadian Clocks in Mood Disorders
    VA I01BX001146Jan 1, 2012 - Sep 30, 2020
    Role: Principal Investigator
    Circadian Clock Cells: Autonomy Persistence and Calcium Dependence
    NIH/NIMH R01MH082945Jun 1, 2008 - Jun 28, 2013
    Role: Principal Investigator
    Circadian Clock Cells: Autonomy, Coupling, and Subtypes
    NIH/NIMH K08MH067657Apr 1, 2003 - Mar 31, 2008
    Role: Principal Investigator

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    Collapse Bibliographic 
    Collapse Publications
    Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Researchers can login to make corrections and additions, or contact us for help.
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    1. McCarthy MJ, Wei H, Nievergelt CM, Stautland A, Maihofer AX, Welsh DK, Shilling P, Alda M, Alliey-Rodriguez N, Anand A, Andreasson OA, Balaraman Y, Berrettini WH, Bertram H, Brennand KJ, Calabrese JR, Calkin CV, Claasen A, Conroy C, Coryell WH, Craig DW, D'Arcangelo N, Demodena A, Djurovic S, Feeder S, Fisher C, Frazier N, Frye MA, Gage FH, Gao K, Garnham J, Gershon ES, Glazer K, Goes F, Goto T, Harrington G, Jakobsen P, Kamali M, Karberg E, Kelly M, Leckband SG, Lohoff F, McInnis MG, Mondimore F, Morken G, Nurnberger JI, Obral S, Oedegaard KJ, Ortiz A, Ritchey M, Ryan K, Schinagle M, Schoeyen H, Schwebel C, Shaw M, Shekhtman T, Slaney C, Stapp E, Szelinger S, Tarwater B, Zandi PP, Kelsoe JR. Chronotype and cellular circadian rhythms predict the clinical response to lithium maintenance treatment in patients with bipolar disorder. Neuropsychopharmacology. 2019 Feb; 44(3):620-628. PMID: 30487653.
      View in: PubMed
    2. Hühne A, Welsh DK, Landgraf D. Prospects for circadian treatment of mood disorders. Ann Med. 2018 Nov 10; 1-18. PMID: 30265156.
      View in: PubMed
    3. Noguchi T, Harrison EM, Sun J, May D, Ng A, Welsh DK, Gorman MR. Circadian rhythm bifurcation induces flexible phase resetting by reducing circadian amplitude. Eur J Neurosci. 2018 Jul 25. PMID: 30044021.
      View in: PubMed
    4. Walton ZE, Patel CH, Brooks RC, Yu Y, Ibrahim-Hashim A, Riddle M, Porcu A, Jiang T, Ecker BL, Tameire F, Koumenis C, Weeraratna AT, Welsh DK, Gillies R, Alwine JC, Zhang L, Powell JD, Dang CV. Acid Suspends the Circadian Clock in Hypoxia through Inhibition of mTOR. Cell. 2018 Jun 28; 174(1):72-87.e32. PMID: 29861175.
      View in: PubMed
    5. Mei L, Fan Y, Lv X, Welsh DK, Zhan C, Zhang EE. Long-term in vivo recording of circadian rhythms in brains of freely moving mice. Proc Natl Acad Sci U S A. 2018 04 17; 115(16):4276-4281. PMID: 29610316.
      View in: PubMed
    6. Porcu A, Riddle M, Dulcis D, Welsh DK. Photoperiod-Induced Neuroplasticity in the Circadian System. Neural Plast. 2018; 2018:5147585. PMID: 29681926.
      View in: PubMed
    7. Noguchi T, Leise TL, Kingsbury NJ, Diemer T, Wang LL, Henson MA, Welsh DK. Calcium Circadian Rhythmicity in the Suprachiasmatic Nucleus: Cell Autonomy and Network Modulation. eNeuro. 2017 Jul-Aug; 4(4). PMID: 28828400.
      View in: PubMed
    8. Diemer T, Landgraf D, Noguchi T, Pan H, Moreno JL, Welsh DK. Cellular circadian oscillators in the suprachiasmatic nucleus remain coupled in the absence of connexin-36. Neuroscience. 2017 08 15; 357:1-11. PMID: 28576728.
      View in: PubMed
    9. Kiessling S, Beaulieu-Laroche L, Blum ID, Landgraf D, Welsh DK, Storch KF, Labrecque N, Cermakian N. Enhancing circadian clock function in cancer cells inhibits tumor growth. BMC Biol. 2017 02 14; 15(1):13. PMID: 28196531.
      View in: PubMed
    10. Beesley S, Noguchi T, Welsh DK. Cardiomyocyte Circadian Oscillations Are Cell-Autonomous, Amplified by ß-Adrenergic Signaling, and Synchronized in Cardiac Ventricle Tissue. PLoS One. 2016; 11(7):e0159618. PMID: 27459195; PMCID: PMC4961434.
    11. Roberts L, Leise TL, Welsh DK, Holmes TC. Functional Contributions of Strong and Weak Cellular Oscillators to Synchrony and Light-shifted Phase Dynamics. J Biol Rhythms. 2016 08; 31(4):337-51. PMID: 27221103.
      View in: PubMed
    12. McCarthy MJ, Wei H, Landgraf D, Le Roux MJ, Welsh DK. Disinhibition of the extracellular-signal-regulated kinase restores the amplification of circadian rhythms by lithium in cells from bipolar disorder patients. Eur Neuropsychopharmacol. 2016 08; 26(8):1310-9. PMID: 27216486.
      View in: PubMed
    13. Landgraf D, Joiner WJ, McCarthy MJ, Kiessling S, Barandas R, Young JW, Cermakian N, Welsh DK. The mood stabilizer valproic acid opposes the effects of dopamine on circadian rhythms. Neuropharmacology. 2016 08; 107:262-270. PMID: 27033596.
      View in: PubMed
    14. Landgraf D, Long JE, Proulx CD, Barandas R, Malinow R, Welsh DK. Genetic Disruption of Circadian Rhythms in the Suprachiasmatic Nucleus Causes Helplessness, Behavioral Despair, and Anxiety-like Behavior in Mice. Biol Psychiatry. 2016 12 01; 80(11):827-835. PMID: 27113500.
      View in: PubMed
    15. Welsh DK, Noguchi T. Erratum: Cellular Bioluminescence Imaging. Cold Spring Harb Protoc. 2016 Mar 01; 2016(3):pdb.err091751. PMID: 26933240.
      View in: PubMed
    16. Noguchi T, Lo K, Diemer T, Welsh DK. Lithium effects on circadian rhythms in fibroblasts and suprachiasmatic nucleus slices from Cry knockout mice. Neurosci Lett. 2016 Apr 21; 619:49-53. PMID: 26930624.
      View in: PubMed
    17. Landgraf D, Wang LL, Diemer T, Welsh DK. NPAS2 Compensates for Loss of CLOCK in Peripheral Circadian Oscillators. PLoS Genet. 2016 Feb; 12(2):e1005882. PMID: 26895328; PMCID: PMC4760943.
    18. Barandas R, Landgraf D, McCarthy MJ, Welsh DK. Circadian Clocks as Modulators of Metabolic Comorbidity in Psychiatric Disorders. Curr Psychiatry Rep. 2015 Dec; 17(12):98. PMID: 26483181.
      View in: PubMed
    19. Landgraf D, Long JE, Welsh DK. Depression-like behaviour in mice is associated with disrupted circadian rhythms in nucleus accumbens and periaqueductal grey. Eur J Neurosci. 2016 05; 43(10):1309-20. PMID: 26414405.
      View in: PubMed
    20. McCarthy MJ, Le Roux MJ, Wei H, Beesley S, Kelsoe JR, Welsh DK. Calcium channel genes associated with bipolar disorder modulate lithium's amplification of circadian rhythms. Neuropharmacology. 2016 Feb; 101:439-48. PMID: 26476274.
      View in: PubMed
    21. Veazey RS, Amedee A, Wang X, Bernice Kaack M, Porretta C, Dufour J, Welsh D, Happel K, Pahar B, Molina PE, Nelson S, Bagby GJ. Chronic Binge Alcohol Administration Increases Intestinal T-Cell Proliferation and Turnover in Rhesus Macaques. Alcohol Clin Exp Res. 2015 Aug; 39(8):1373-9. PMID: 26146859; PMCID: PMC4515194.
    22. Kripke DF, Elliott JA, Welsh DK, Youngstedt SD. Photoperiodic and circadian bifurcation theories of depression and mania. F1000Res. 2015; 4:107. PMID: 26180634; PMCID: PMC4490783.
    23. Landgraf D, Long J, Der-Avakian A, Streets M, Welsh DK. Dissociation of learned helplessness and fear conditioning in mice: a mouse model of depression. PLoS One. 2015; 10(4):e0125892. PMID: 25928892; PMCID: PMC4416012.
    24. Roberts L, Leise TL, Noguchi T, Galschiodt AM, Houl JH, Welsh DK, Holmes TC. Light evokes rapid circadian network oscillator desynchrony followed by gradual phase retuning of synchrony. Curr Biol. 2015 Mar 30; 25(7):858-67. PMID: 25754644; PMCID: PMC4399721.
    25. Landgraf D, McCarthy MJ, Welsh DK. Circadian clock and stress interactions in the molecular biology of psychiatric disorders. Curr Psychiatry Rep. 2014 Oct; 16(10):483. PMID: 25135782.
      View in: PubMed
    26. Ramanathan C, Xu H, Khan SK, Shen Y, Gitis PJ, Welsh DK, Hogenesch JB, Liu AC. Cell type-specific functions of period genes revealed by novel adipocyte and hepatocyte circadian clock models. PLoS Genet. 2014 Apr; 10(4):e1004244. PMID: 24699442; PMCID: PMC3974647.
    27. Landgraf D, McCarthy MJ, Welsh DK. The role of the circadian clock in animal models of mood disorders. Behav Neurosci. 2014 Jun; 128(3):344-59. PMID: 24660657.
      View in: PubMed
    28. Noguchi T, Wang LL, Welsh DK. Fibroblast PER2 circadian rhythmicity depends on cell density. J Biol Rhythms. 2013 Jun; 28(3):183-92. PMID: 23735497; PMCID: PMC3760388.
    29. McCarthy MJ, Fernandes M, Kranzler HR, Covault JM, Welsh DK. Circadian clock period inversely correlates with illness severity in cells from patients with alcohol use disorders. Alcohol Clin Exp Res. 2013 Aug; 37(8):1304-10. PMID: 23550834; PMCID: PMC3706548.
    30. Evans JA, Pan H, Liu AC, Welsh DK. Cry1-/- circadian rhythmicity depends on SCN intercellular coupling. J Biol Rhythms. 2012 Dec; 27(6):443-52. PMID: 23223370; PMCID: PMC3578226.
    31. McCarthy MJ, Welsh DK. Cellular circadian clocks in mood disorders. J Biol Rhythms. 2012 Oct; 27(5):339-52. PMID: 23010657.
      View in: PubMed
    32. Cohen AL, Leise TL, Welsh DK. Bayesian statistical analysis of circadian oscillations in fibroblasts. J Theor Biol. 2012 Dec 07; 314:182-91. PMID: 22982138; PMCID: PMC3478438.
    33. Welsh DK, Noguchi T. Cellular bioluminescence imaging. Cold Spring Harb Protoc. 2012 Aug 01; 2012(8). PMID: 22854570.
      View in: PubMed
    34. Hirota T, Lee JW, St John PC, Sawa M, Iwaisako K, Noguchi T, Pongsawakul PY, Sonntag T, Welsh DK, Brenner DA, Doyle FJ, Schultz PG, Kay SA. Identification of small molecule activators of cryptochrome. Science. 2012 Aug 31; 337(6098):1094-7. PMID: 22798407; PMCID: PMC3589997.
    35. Noguchi T, Wang CW, Pan H, Welsh DK. Fibroblast circadian rhythms of PER2 expression depend on membrane potential and intracellular calcium. Chronobiol Int. 2012 Jul; 29(6):653-64. PMID: 22734566; PMCID: PMC3491983.
    36. Leise TL, Wang CW, Gitis PJ, Welsh DK. Persistent cell-autonomous circadian oscillations in fibroblasts revealed by six-week single-cell imaging of PER2::LUC bioluminescence. PLoS One. 2012; 7(3):e33334. PMID: 22479387; PMCID: PMC3315561.
    37. McCarthy MJ, Nievergelt CM, Kelsoe JR, Welsh DK. A survey of genomic studies supports association of circadian clock genes with bipolar disorder spectrum illnesses and lithium response. PLoS One. 2012; 7(2):e32091. PMID: 22384149; PMCID: PMC3285204.
    38. Meeker K, Harang R, Webb AB, Welsh DK, Doyle FJ, Bonnet G, Herzog ED, Petzold LR. Wavelet measurement suggests cause of period instability in mammalian circadian neurons. J Biol Rhythms. 2011 Aug; 26(4):353-62. PMID: 21775294.
      View in: PubMed
    39. Foley NC, Tong TY, Foley D, Lesauter J, Welsh DK, Silver R. Characterization of orderly spatiotemporal patterns of clock gene activation in mammalian suprachiasmatic nucleus. Eur J Neurosci. 2011 May; 33(10):1851-65. PMID: 21488990; PMCID: PMC3423955.
    40. 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. Duplications of the neuropeptide receptor gene VIPR2 confer significant risk for schizophrenia. Nature. 2011 Mar 24; 471(7339):499-503. PMID: 21346763; PMCID: PMC3351382.
    41. Ko CH, Yamada YR, Welsh DK, Buhr ED, Liu AC, Zhang EE, Ralph MR, Kay SA, Forger DB, Takahashi JS. Emergence of noise-induced oscillations in the central circadian pacemaker. PLoS Biol. 2010 Oct 12; 8(10):e1000513. PMID: 20967239; PMCID: PMC2953532.
    42. Welsh DK, Takahashi JS, Kay SA. Suprachiasmatic nucleus: cell autonomy and network properties. Annu Rev Physiol. 2010; 72:551-77. PMID: 20148688; PMCID: PMC3758475.
    43. Westermark PO, Welsh DK, Okamura H, Herzel H. Quantification of circadian rhythms in single cells. PLoS Comput Biol. 2009 Nov; 5(11):e1000580. PMID: 19956762; PMCID: PMC2776301.
    44. Mirsky HP, Liu AC, Welsh DK, Kay SA, Doyle FJ. A model of the cell-autonomous mammalian circadian clock. Proc Natl Acad Sci U S A. 2009 Jul 07; 106(27):11107-12. PMID: 19549830; PMCID: PMC2699375.
    45. Wang LM, Dragich JM, Kudo T, Odom IH, Welsh DK, O'Dell TJ, Colwell CS. Expression of the circadian clock gene Period2 in the hippocampus: possible implications for synaptic plasticity and learned behaviour. ASN Neuro. 2009 Jun 10; 1(3). PMID: 19570032; PMCID: PMC2695588.
    46. Liu AC, Tran HG, Zhang EE, Priest AA, Welsh DK, Kay SA. Redundant function of REV-ERBalpha and beta and non-essential role for Bmal1 cycling in transcriptional regulation of intracellular circadian rhythms. PLoS Genet. 2008 Feb 29; 4(2):e1000023. PMID: 18454201; PMCID: PMC2265523.
    47. Pulivarthy SR, Tanaka N, Welsh DK, De Haro L, Verma IM, Panda S. Reciprocity between phase shifts and amplitude changes in the mammalian circadian clock. Proc Natl Acad Sci U S A. 2007 Dec 18; 104(51):20356-61. PMID: 18077393; PMCID: PMC2154435.
    48. Forger D, Gonze D, Virshup D, Welsh DK. Beyond intuitive modeling: combining biophysical models with innovative experiments to move the circadian clock field forward. J Biol Rhythms. 2007 Jun; 22(3):200-10. PMID: 17517910.
      View in: PubMed
    49. Liu AC, Welsh DK, Ko CH, Tran HG, Zhang EE, Priest AA, Buhr ED, Singer O, Meeker K, Verma IM, Doyle FJ, Takahashi JS, Kay SA. Intercellular coupling confers robustness against mutations in the SCN circadian clock network. Cell. 2007 May 04; 129(3):605-16. PMID: 17482552; PMCID: PMC3749832.
    50. Welsh DK. Gate cells see the light. J Biol Rhythms. 2007 Feb; 22(1):26-8. PMID: 17229922.
      View in: PubMed
    51. Welsh DK. VIP activates and couples clock cells. Focus on "Disrupted neuronal activity rhythms in the suprachiasmatic nucleus of vasoactive intestinal polypeptide-deficient mice". J Neurophysiol. 2007 Mar; 97(3):1885-6. PMID: 17251373.
      View in: PubMed
    52. Sato TK, Yamada RG, Ukai H, Baggs JE, Miraglia LJ, Kobayashi TJ, Welsh DK, Kay SA, Ueda HR, Hogenesch JB. Feedback repression is required for mammalian circadian clock function. Nat Genet. 2006 Mar; 38(3):312-9. PMID: 16474406; PMCID: PMC1994933.
    53. Welsh DK, Kay SA. Bioluminescence imaging in living organisms. Curr Opin Biotechnol. 2005 Feb; 16(1):73-8. PMID: 15722018.
      View in: PubMed
    54. Welsh DK, Imaizumi T, Kay SA. Real-time reporting of circadian-regulated gene expression by luciferase imaging in plants and mammalian cells. Methods Enzymol. 2005; 393:269-88. PMID: 15817294.
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    55. Welsh DK, Yoo SH, Liu AC, Takahashi JS, Kay SA. Bioluminescence imaging of individual fibroblasts reveals persistent, independently phased circadian rhythms of clock gene expression. Curr Biol. 2004 Dec 29; 14(24):2289-95. PMID: 15620658; PMCID: PMC3777438.
    56. Monk TH, Welsh DK. The role of chronobiology in sleep disorders medicine. Sleep Med Rev. 2003 Dec; 7(6):455-73. PMID: 15018090.
      View in: PubMed