Altered Calmodulin Response to Light in the Suprachiasmatic
Nucleus of PAC
1
Receptor Knockout Mice Revealed
by Proteomic Analysis
Jan Fahrenkrug,*
,1
Jens Hannibal,
1
Bent Honoré,
2
and Henrik Vorum
2
1
Department of Clinical Biochemistry, Bispebjerg Hospital, University of Copenhagen,
DK-2400 Copenhagen NV, Denmark; and
2
Institute of Medical Biochemistry, University
of Aarhus, DK-8000 Aarhus C, Denmark
Received July 2, 2004; Accepted July 29, 2004
Abstract
In mammals circadian rhythms are generated by a light-entrainable oscillator located in the hypothalamic
suprachiasmatic nucleus (SCN). Light signals reach the SCN via a monosynaptic neuronal pathway, the retino-
hypothalamic tract, originating in a subset of retinal ganglion cells. The nerve terminals of these cells contain
the classical neurotransmitter glutamate and the neuropeptide pituitary adenylate cyclase-activating polypep-
tide (PACAP), and there is evidence that these two transmitters interact to mediate photoentrainment of the
oscillator in the SCN. To elucidate light-provoked PACAPreceptor signaling we used proteomic analysis. Wild-
type mice and mice lacking the PAC
1
receptor (PAC
1
–/–
) were light stimulated at early night, and the SCN was
examined for proteins that were differentially expressed using two-dimensional gel electrophoresis and iden-
tification by tandem mass spectrometry. The most striking finding, which was subsequently confirmed by West-
ern blotting, was a significant reduction of calmodulin (CaM) in wild-type mice as compared with PAC
1
–/–
mice.
Analysis at the mRNA level by quantitative in situ hybridization histochemistry was inconclusive, indicating
that a translational mechanism might be involved. The findings indicate that PAC
1
receptor signaling in the
SCN in response to light stimulation induces a down-regulation of CaM expression and that CaM is involved
in the photic-entrainment mechanism.
Index Entries: Calmodulin; two-dimensional gel electrophoresis; knockout; PAC
1
receptor; photic entrain-
ment; pituitary adenylate cyclase-activating polypeptide; proteomics; suprachiasmatic nucleus.
Introduction
In mammals daily physiological and behavioral
rhythms are governed by a biological clock located
in the suprachiasmatic nucleus (SCN) of the hypo-
thalamus (Klein et al., 1991). The molecular machin-
ery driving the brain’s biological clock consists of a
number of clock genes, which, in double autoregu-
latory feedback loops, interact and regulate their own
transcription within the individual SCN neurons
(Reppert and Weaver, 2002). To maintain synchrony
with the solar day/night cycle, the clock is adjusted
daily by light, a process called photoentrainment
(Roenneberg and Foster, 1997); but the mechanisms
by which external light cues modify the activity of
the pacemaker in the SCN are not completely under-
stood. The major input pathway mediating infor-
mation on environmental light and darkness
originates in a subpopulation of retinal ganglion cells
that project to the SCN via the retinohypothalamic
Journal of Molecular Neuroscience
Copyright © 2005 Humana Press Inc.
All rights of any nature whatsoever reserved.
ISSN0895-8696/02/25:251–258/$30.00
Journal of Molecular Neuroscience 251 Volume 25, 2005
*Author to whom all correspondence and reprint request should be addressed. E-mail: bbhjanf@inet.uni2.dk
ORIGINAL ARTICLE