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Neuroscience Letters206 (1996) 97-100
NHllOSCHCI
LHTEIIS
Nerve growth factor phase shifts circadian activity rhythms
in Syrian hamsters
Keshavan G, Bina a, Benjamin Rusaka,b, *
aDepartmentof Psychology, Dalhousie University, Halifax, Nova Scotia, B3H 4Jl, Canada
bDepartment of Pharmacology, Dalhousie University, Halifax, Nova Scotia, B3H 4H7, Canada
Received29 December1995;revisedversionreceived 2 February1996;accepted2 February1996
Abstract
Nerve growth factor (NGF) receptors are found in high density in the rodent suprachiasmatic nucleus (SCN), a site which regulates
mammalian circadian rh)~hms. We examined the effects of NGF (40 ng) or vehicle injections into the SCN, at circadian times (CT) 6,
14 or 22 on activity rhythms in hamsters maintained in constant darkness. NGF caused phase advances at CT6 (30.9 rain) and CT22
(36.9 min), and phase delays at CT14 (31.2 rain). Saline and cytochrome-c administration had no phase-shifting effects at CT6 and
CT22, but at CT14 cytochrome-c produced large phase delays, implying that NGF-induced delays at this phase may be non-specific.
Similarities between NGF-induced shifts and those elicited by the cholinergic agonist carbachol suggest a common mode of action.
Keywords: Suprachiasmatic; Entrainment; Nerve growth factor; Phase response curve; Cholinergic; Acetylcholine
The mammalian suprachiasmatic nucleus (SCN) is
responsible for the generation of circadian rhythms and
their synchronization (entrainment) to lighting cycles [9].
Acetylcholine (ACh) has been proposed to be involved in
entrainment because iniections of carbachol, a cholinergic
agonist, intraventricularly or into the SCN, phase shift
activity rhythms in rodents, somewhat as light does [19].
These effects of carbachol may involve presynaptic
modulation of the release of an excitatory amino acid [6].
Receptors for nerve growth factor (NGF; NGF-R) are
synthesized in choline, rgic neurons and bind NGF re-
leased by their postsynaptic target neurons, forming a
NGF/NGF-R complex that exerts a biological effect criti-
cal to the survival and maintenance of these neurons [24].
The SCN contains high levels of NGF-R immunoreactiv-
ity [22], at least some of which is located on terminals of
basal forebrain cholinergic neurons and retinal ganglion
cells projecting to the SCN [3]. In addition, neurons in the
SCN have been shown to contain both NGF [2] and its
precursor [21], We therefore examined the effects of NGF
applications into the SCN on circadian rhythms. A pre-
liminary report of this study has been presented [2].
* Correspondingauthor. Tell.: +1 902 4942159; fax: +1 902 4942159;
e-mail: rusak@ac.dal.ca.
Fifty adult male hamsters (Mesocricetus auratus;
LVG:lak, 80-100 g; Charles River, St. Constant, Qu6bec)
were implanted under pentobarbital anesthesia (80 mg/kg)
with stainless steel guide cannulas (22 gauge; Plastics
One, Roanoke, VA) directed stereotaxically at the SCN
(0.5 mm anterior to bregma, 0.3 mm lateral to midline,
and 8.4 mm ventral to the skull surface, with the incisor
bar 2 mm below the interaural line). After recovery from
anesthesia, animals were housed individually in cages
equipped with activity wheels and maintained under con-
stant dim red light (DD). Wheel-running activity was
monitored continuously on an event recorder (Esterline-
Angus, Indianapolis) and activity counts were recorded
every 10 min using an Apple computer. Event recorder
charts were used to measure activity rhythm periods and
phase shifts, while the actograms presented in the figures
were plotted from computer files.
After a stable free-running rhythm was established,
hamsters were injected intracranially with 2/tl of one of
the following: 0.9% saline, a saline solution of NGF
(20 ng//~l saline; Sigma Chemical Co.), or a solution of
cytochrome-c (CYT; 300 ng//tl; Sigma Chemical Co.),
which is physicochemically similar to NGF, but lacks its
biological activity [1]. Each injection was made over 2-
3 min using a Hamilton syringe connected via plastic
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