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1 Dept. of Biology, University of Virginia, Charlottesville, Virginia, United States
* To whom correspondence should be addressed. E-mail: mm7e{at}virginia.edu.
The circadian clock in the suprachiasmatic nucleus (SCN) maintains phase synchrony among circadian oscillators throughout the organism. Light signals entrain the SCN but timed, limited meal access acts as an overriding time cue for several peripheral tissues. In one peripheral oscillator, the submaxillary salivary glands, temporal restriction of meals fails to entrain gene expression. In day-fed rats, submaxillary gland rhythms in expression of the clock gene Period1 either stay entrained to the light cycle (peaking at night) or become arrhythmic. This suggests that feeding cues compete weakly with light cycle cues to set the phase of clock genes in this tissue. Since the submaxillary glands receive sympathetic innervation originating in the SCN, we attempted to assess the role of this neural input in phase control of submaxillary Period1 expression. To do this, we sympathetically denervated the submaxillary glands before subjecting rats to daytime restricted feeding. Following denervation Period1 rhythms in all submaxillary glands shifted phase 180° and entrained to daytime feeding. These results support the hypothesis that peripheral oscillators may receive multiple signals contributing to their phase of entrainment. Sympathetic efferents from the SCN can relay light cycle information while other external cues may reach tissues through other efferents or non-neural pathways. In an abnormal, disruptive regime such as daytime restricted feeding these different signals compete. Arrhythmicity may result if one signal is not clearly dominant. Elimination of the dominant signal (e.g. surgical sympathectomy) may allow a secondary signal to control phase.
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