DAVID K. WELSH, M.D., Ph.D.

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DAVID K. WELSH, M.D., Ph.D.

Department of Psychiatry

University of California, San Diego

9500 Gilman Drive #0603, La Jolla, CA 92093
-
0603

e
-
mail:
welshdk@ucsd.edu
; phone: (858) 246
-
0894


Academic Positions:

Universi
ty of California, San Diego, Department of Psychiatry


Assistant Clinical Professor, 4/03
-

6/03


Assistant Professor, 7/03
-

date


NIMH Mentored Clinical Scientist Development Award, K08 MH067657, 4/03
-

3/08


NIMH Research Project Grant, R01 MH082945, 3/
08
-

date

Veterans Affairs San Diego Healthcare System, San Diego, CA, Staff Physician, 7/03
-

date


Career Development Award 2, 11/08
-

date


Education:

Stanford University, Stanford, CA, 9/79
-

6/84, B.A.S. in Biological Sciences and Psychology,
With Dis
tinction, Phi Beta Kappa

Harvard Medical School, Boston, MA, 9/85
-

6/97, M.D., Ph.D. in Neurobiology


Predoctoral Fellow, Howard Hughes Medical Institute, 7/88
-

6/93


Chronobiology Research Program, U.S. Air Force Office of Scientific Research, 7/93
-

8/
95


Medical Scientist Training Program, National Institutes of Health, 7/93
-

6/97

Marine Biological Laboratory, Woods Hole, MA, 6/89
-

8/89, Neurobiology course

University Health Center of Pittsburgh, Western Psychiatric Institute and Clinic, 6/97
-

6/98

Internship in Medicine and General Psychiatry

Western Psychiatric Institute & Clinic, Pittsburgh, 7/98
-

6/01, Residency in General Psychiatry



Outstanding Resident Award, National Institute of Mental Health, 1999


Laughlin Fellow, American College of Psy
chiatrists, 2000


Lilly Fellow, Society of Biological Psychiatry, 2000

University of California, San Diego, 7/01
-

3/03, Research Fellow, Department of Psychiatry


Postdoctoral Fellowship in Biological Psychiatry and Neuroscience, UCSD


Research Associate,

Department of Cell Biology, The Scripps Research Institute


Oral Presentations at National or International Meetings:

Gerontological Society of America, San Francisco, 1983

Society for Neuroscience, New Orleans, 1991

Society for Neuroscience, Miami, 1994

Gordon Research Conference on Chronobiology, Il Ciocco, Barga, Italy, 1995

Society for Research on Biological Rhythms, Jacksonville, FL, 1996

Suprachiasmatic Nucleus 25th Anniversary Symposium, Harvard Medical School, Boston, 1997

Workshop on Biomathematic
al Models of Circadian Rhythmicity, Sleep Regulation, and
Neurobehavioral Function in Humans, Boston, 1999

Society for Research on Biological Rhythms, Whistler, BC, Canada, 2004

Society for Research on Biological Rhythms, Destin, FL, 2006

Workshop on Model
ing Circadian Rhythmicity, Sleep Regulation, and Neurobehavioral
Function, Joint SIAM/SMB Conference on the Life Sciences, Raleigh, NC, 2006

David K. Welsh

2

12/4/13

Imaging Neurons and Neural Activity, Cold Spring Harbor Laboratory, NY, 2007

Gordon Research Conference on Chronobi
ology, Aussois, France, 2007

Physiological Society of Japan, Tokyo, 2008

Society for Research on Biological Rhythms, Destin, FL, 2008

International Congress of Physiological Sciences, Kyoto, Japan, 2009

International Symposium on Biological Rhythm, Sapporo
, Japan, 2009

Society for Research on Biological Rhythms, Destin, FL, 2010

Assembling a Multi
-
Cellular Circadian Pacemaker, Lorentz Center Workshop, Leiden
University, Leiden, The Netherlands, 2010

“It’s About Time!”


Biological Clocks and Neural Function
, Rushton Lecture Series, Florida
State University, Tallahassee, FL, 2011


Publications
:


1) Wandell BA, Welsh D, Maloney L. Adaptation in the long
-
wavelength pathways.
Vision

Res

22:1071
-
1074, 1982.


2) Wandell BA, Ahumada P, Welsh D. Reaction times to we
ak test lights.
Vision Res

24:647
-
652, 1984.


3) Welsh DK, Richardson GS, Dement WC. A circadian rhythm of hippocampal theta activity in
the mouse.
Physiol Behav

35:533
-
538, 1985.


4) Welsh DK, Nino
-
Murcia G, Gander PH, Keenan S, Dement WC. Regular 48
-
hour

cycling of
sleep duration and mood in a 35
-
year
-
old woman: use of lithium in time isolation.
Biol
Psychiatry

21:527
-
537, 1986.


5) Welsh DK, Engel EMRA, Richardson GS, Dement WC. Precision of circadian wake and
activity onset timing in the mouse.
J Comp P
hysiol A

158:827
-
834, 1986.


6) Welsh DK, Richardson GS, Dement WC. Effect of age on the circadian pattern of sleep and
wakefulness in the mouse.
J Gerontology

41:579
-
586, 1986.


7) Welsh DK, Richardson GS, Dement WC. Effect of running wheel availability o
n circadian
patterns of sleep and wakefulness in mice.
Physiol Behav

43:771
-
777, 1988.


8) Nadakavukaren JJ, Welsh DK, Reppert SM. Aluminum fluoride reveals a phosphoinositide
system within the suprachiasmatic region of rat hypothalamus.
Brain Res

507:181
-
188, 1990.


9) Welsh DK, Moore
-
Ede MC. Lithium lengthens circadian period in a diurnal primate,
Saimiri
sciureus
.
Biol Psychiatry

28:117
-
126, 1990.


10) Welsh DK, Logothetis DE, Meister M, Reppert SM. Individual neurons dissociated from rat
suprachiasmatic

nucleus express independently phased circadian firing rhythms.
Neuron

14:697
-
706, 1995.


David K. Welsh

3

12/4/13

11) Welsh DK, Reppert SM. Gap junctions couple astrocytes but not neurons in dissociated
cultures of rat suprachiasmatic nucleus.
Brain Res

706:30
-
36, 1996.


12) Noll

E, Welsh DK, Fan M
-
H, Canales M, Black PM. Towards a lineage
-
based classification
system for gliomas: innovation or imagination?
Crit Rev Neurosurg

8:6
-
18, 1998.


13) Monk TH, Buysse DJ, Welsh DK, Kennedy KS, Rose LR. A sleep diary and questionnaire
study

of naturally short sleepers.
J Sleep Res

10:173
-
179, 2001.


14) Monk TH, Welsh DK. The role of chronobiology in sleep disorders medicine.
Sleep Med Rev
7:455
-
473, 2003.


15) Welsh DK, Yoo SH, Liu AC, Takahashi JS, Kay SA. Bioluminescence imaging of indivi
dual
fibroblasts reveals persistent, independently phased circadian rhythms of clock gene
expression.
Curr Biol

14:2289
-
2295, 2004.


16) Welsh DK, Kay SA. Bioluminescence imaging in living organisms.
Curr Opin Biotechnol

16:73
-
78, 2005.


17) Welsh DK, Ima
izumi T, Kay SA. Real
-
time reporting of circadian
-
regulated gene expression
by luciferase imaging in plants and mammalian cells.
Meth Enzymol

393:269
-
288, 2005.


18) Welsh DK. Genetics of circadian rhythm disorders.
Psychiatric Times

22(14):37
-
40, 2005.


1
9) 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

38:312
-
319, 2006. [
PMCID: PMC1994933
]


20) Welsh DK. Comment: Gat
e cells see the light.
J Biol Rhythms

22:26
-
28, 2007.


21) 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

97
:1885
-
1886, 2007.


22) Liu AC*, Welsh DK*, Ko CH, Tran HG, Zhang EE, Priest AA, Buhr ED, Singer O, Meeker
K, Verma IM, Doyle FJ III, Takahashi JS, Kay SA. Intercellular coupling confers robustness
against mutations in the SCN circadian clock network.
Cell

129:605
-
616, 2007.


23) 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

22:200
-
210, 2007.


24) 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 USA

104:20356
-
20361, 2007. [
PMCID: PMC2154435]


David K. Welsh

4

12/4/13

25) Yan L, Karatsoreos I, LeSauter J, Welsh DK, Kay S, Fole
y D, Silver R. Exploring
spatiotemporal organization of SCN circuits.
Cold Spring Harb Symp Quant Biol

72:527
-
541, 2007.


26) Liu AC, Tran HG, Zhang EE, Priest AA, Welsh DK, Kay SA. Redundant function of REV
-
ERB


and


and non
-
essential role for Bmal1 cycl
ing in transcriptional regulation of
intracellular circadian rhythms.
PLoS Genet

4(2):e1000023, 2008. [
PMCID: PMC2265523
]


27) Mirsky HP, Liu AC, Welsh DK, Kay SA, Doyle FJ 3
rd
. A model of the cell
-
autonomous
mammalian circadian clock.
Proc Natl Acad Sci U
SA

106:11107
-
12, 2009. [
PMCID:
PMC2699375
]


28) Wang LM
-
C, 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

1(3):art:e00012, 2009. [
PMCID:
PMC2695588
]


29) Westermark PO, Welsh DK, Okamura H, Herzel H. Quantification of circadian rhythms in
single cells.
PLoS Comput Biol

5(11):e1000580, 2009. [
PMCID: PMC2776301
]


30) Welsh DK. Limits of single
-
cell au
tonomy in the suprachiasmatic nucleus.
Sleep Biol
Rhythms

7:252
-
9, 2009. [PMC Exempt]


31) Welsh DK, Ptácek LJ. Circadian rhythm dysregulation in the elderly: advanced sleep phase
syndrome. In: Pandi
-
Perumal SR, Monti JM, Monjan AA (Eds),
Principles and Pr
actice of
Geriatric Sleep Medicine
, Cambridge, UK: Cambridge University Press, 2010, pp. 131
-
142.
[PMC Exempt]


32) Welsh DK, Takahashi JS, Kay SA. Suprachiasmatic nucleus: cell autonomy and network
properties.
Annu Rev Physiol

72:551
-
77, 2010. [PMC Exempt
]


33) 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

8(10):e1000513, 2010
. [
PMCID: PMC2953532]


34) Welsh DK, Noguc
hi T. Cellular bioluminescence imaging. In: Yuste R (Ed),
Imaging: A
Laboratory Manual
, Cold Spring Harbor, NY: Cold Spring Harbor Laboratory Press, 2011,
pp. 369
-
385
. [PMC Exempt]


35) 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, Aquila MD, Morris DW, Gill M,
Corvin A, Insel PA, McClellan J, King MC, Karayiourgou M, Levy DL, DeLisi LE,
Sebat J.
Genomic duplications of the neuropeptide receptor VIPR2 confer significant risk for
schizophrenia.
Nature

471:499
-
503, 2011
.


David K. Welsh

5

12/4/13

36) Foley N, Tong TY, Foley D, LeSauter J, Welsh DK, Silver R. Characterization of orderly
spatiotemporal patterns of clo
ck gene activation in mammalian suprachiasmatic nucleus.
Eur J Neurosci

33:1851
-
65, 2011.


37)
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 circadi
an neurons
.

J Biol Rhythms

26: 353
-
62, 2011.


38) McCarthy MJ, Nievergelt CM, Shekhtman T, Kripke DF, Welsh DK, Kelsoe JR. Functional
genetic variation in the Rev
-
Erb


pathway and lithium response in the treatment of bipolar
disorder.
Genes Brain Behav
: In

Press, 2011.



* equal contributions