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No. of Pages 6
29 July 2015
Please cite this article in press as: van der Vinne V et al. Food reward without a timing component does not alter the timing of activity under positive
energy balance. Neuroscience (2015), http://dx.doi.org/10.1016/j.neuroscience.2015.07.061
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AbstractCircadian clocks drive daily rhythms in physiology and behavior which allow organisms to anticipate
predictable daily changes in the environment. In most mammals, circadian rhythms result in nocturnal activity patterns
although plasticity of the circadian system allows activity
patterns to shift to dierent times of day. Such plasticity is
seen when food access is restricted to a few hours during
the resting (light) phase resulting in food anticipatory activity
(FAA) in the hours preceding food availability. The mechanisms underlying FAA are unknown but data suggest the
involvement of the reward system and homeostatic regulation of metabolism. We previously demonstrated the isolated
eect of metabolism by inducing diurnality in response to
energetic challenges. Here the importance of reward timing
in inducing daytime activity is assessed. The daily activity
distribution of mice earning palatable chocolate at their preferred time by working in a running wheel was compared with
that of mice receiving a timed palatable meal at noon. Mice
working for chocolate (WFC) without being energetically
challenged increased their total daily activity but this did
not result in a shift to diurnality. Providing a chocolate meal
at noon each day increased daytime activity, identifying food
timing as a factor capable of altering the daily distribution of
activity and rest. These results show that timing of food
reward and energetic challenges are both independently sufcient to induce diurnality in nocturnal mammals. FAA
observed following timed food restriction is likely the result
of an additive eect of distinct regulatory pathways activated
by energetic challenges and food reward. 2015 Published
by Elsevier Ltd. on behalf of IBRO.
*Corresponding author.
E-mail address: vincentvandervinne@gmail.com (V. van der Vinne).
INTRODUCTION
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EXPERIMENTAL PROCEDURES
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Experiment 1: WFC
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The WFC protocol allowed twelve ad libitum fed, vemonth-old mice to additionally earn 45-mg chocolate
pellets (Dustless Precision Pellets, Sucrose, Chocolate
Flavor) by running in a wheel. The workload, the number
of revolutions needed to obtain a chocolate pellet, started
at 500 revs/pellet on day nine and was increased in steps
of 20 revs/pellet daily to 700 revs/pellet, where it was kept
stable (resulting in 37.3 3.5 chocolate pellets per day).
This workload was chosen because it resulted in a
chocolate intake that was approximately twice the intake
observed when chocolate was provided at a xed time,
ensuring that the lack of a phase shift during WFC was
not a result of lowered chocolate intake. The timing of the
chocolate reward in the WFC protocol was computer
controlled and the mice thus received their food reward
each time they reached the workload threshold. Reward
timing thus depended solely on the spontaneous activity
rhythm of the mice. After the WFC protocol (day 26 until
the end), each mouse was provided with 20 pellets daily
at the middle of the light phase (ExT 12) to assess the
eect of timed chocolate feeding (TCF) on the activity
rhythm. Daily checks were performed around one hour
before lights o, to conrm that all chocolate pellets had
been consumed. The daily distribution of activity was
compared for the last 5 days preceding WFC (AL), the
last 5 days of WFC (WFC) and the last 5 days of timed
chocolate feeding.
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Experiment 2: TCF
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Please cite this article in press as: van der Vinne V et al. Food reward without a timing component does not alter the timing of activity under positive
energy balance. Neuroscience (2015), http://dx.doi.org/10.1016/j.neuroscience.2015.07.061
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RESULTS
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Experiment 1: WFC
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Fig. 1. Mice remain nocturnal during working for chocolate (WFC) but diurnality is induced by timed chocolate feeding during the light phase (TCF).
(A) Representative actogram of ad libitum fed mouse undergoing WFC (day 925, hatched area) followed by TCF (day 2639, vertical line). The
daily proportion of activity occurring during the light phase is plotted in the right hand graph. (B) The proportion of activity occurring during the light
phase is unaected by WFC (day 2125) but increases signicantly during TCF (day 3539), compared to ad libitum (AL; day 48) fed mice. (C)
Total daily activity is increased during the last ve days of WFC compared to AL and TCF. *p < 0.05.
Fig. 2. Activity is phase advanced by TCF but not by WFC compared to AL. (A) Average relative activity proles during the last 5 days of AL, WFC
and TCF in 10-min bins with SEM. TCF induced a phase advance of activity. Activity was increased during the middle and late light phase and
decreased during the middle dark phase during TCF compared to AL and WFC. (B) Activity onset, center of gravity (COG) and oset were all phase
advanced by TCF but not by WFC.
Please cite this article in press as: van der Vinne V et al. Food reward without a timing component does not alter the timing of activity under positive
energy balance. Neuroscience (2015), http://dx.doi.org/10.1016/j.neuroscience.2015.07.061
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Fig. 3. Representative actograms showing running wheel and feeding place activity of a mouse provided with chocolate (days 928, 3140) or
additional chow (days 2930, 4160) for two hours during the light phase. Chocolate feeding during the day advanced the main activity bout but did
not induce food anticipatory activity in either running wheel or feeding place activity. The daily proportion of total activity occurring during the light
phase is plotted to the right of each actogram. The total number of consumed chocolate (purple/black) and chow (green/gray) pellets eaten during
the 2-h interval is shown in the middle graph. (For interpretation of the references to color in this gure legend, the reader is referred to the web
version of this article.)
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Experiment 2: TCF
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Please cite this article in press as: van der Vinne V et al. Food reward without a timing component does not alter the timing of activity under positive
energy balance. Neuroscience (2015), http://dx.doi.org/10.1016/j.neuroscience.2015.07.061
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DISCUSSION
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Please cite this article in press as: van der Vinne V et al. Food reward without a timing component does not alter the timing of activity under positive
energy balance. Neuroscience (2015), http://dx.doi.org/10.1016/j.neuroscience.2015.07.061
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REFERENCES
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Please cite this article in press as: van der Vinne V et al. Food reward without a timing component does not alter the timing of activity under positive
energy balance. Neuroscience (2015), http://dx.doi.org/10.1016/j.neuroscience.2015.07.061
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