First published online May 21, 2007
Journal of Experimental Biology 210, 2013-2024 (2007)
Published by The Company of Biologists 2007
doi: 10.1242/jeb.001974
Behavioural and physiological responses to increased foraging effort in male mice
Lobke M. Vaanholt1,*,
Berber De Jong1,
Theodore Garland, Jr2,
Serge Daan1 and
G. Henk Visser1,3
1 University of Groningen, Department of Behavioural Biology, Kerklaan 30,
9751 NN Haren, The Netherlands
2 University of California, Riverside, Department of Biology, Riverside, CA
92521, USA
3 University of Groningen, Centre for Isotope Research, Nijenborgh 4, 9747
AG Groningen, The Netherlands

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Fig. 1. Relationship between spontaneous wheel-running activity (RWA BL) and
maximum wheel-running activity (RWA MX) in control mice (C, open circles) and
mice selectively bred for high wheel-running activity (S, closed circles).
Linear regression gave the following equations: combining both groups, RWA
MX=0.35 RWA BL+20.1 (r2=0.09, n.s.); for C mice, RWA
MX=0.22 RWA BL+27.6 (r2=0.02, n.s.) and for S mice,
RWA MX=0.84 RWA BL+15.2 (r2=0.47, n.s.). The line shows
where x=y.
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Fig. 2. Circadian pattern of wheel-running activity in control (C, open symbols)
and activity-selected mice (S, closed symbols) running spontaneously (circles)
or running for food (triangles). Each symbol plots the mean distance ran in
hourly bins (e.g. bin 12=from 12:00 until 13:00). Vertical bars are
inter-individual s.e.m. The black bar on top represents the dark phase.
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Fig. 3. Development of wheel-running activity, body mass and food intake during
training and at a workload of 90% from the maximal capacity in Workload
animals (C and S groups pooled). Spontaneous wheel-running activity is shown
for the 2 weeks prior to the training period (day 0). Circles show the
development of the different variables during the experiment in the Workload
animals, and triangles represent mean values for mice in the Ad-lib
group.
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Fig. 4. Relationship between body mass and metabolic rates (A) and between
wheel-running activity and metabolic rates (B) at baseline (open symbols) and
workload (closed symbols) conditions in Workload animals. Triangles represent
the RMR and circles represent DEE. Regression lines for all relationships are
drawn. For equations of the regression lines, r2 and
P values, see Table 4.
Results of multiple regressions are presented in the text.
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Fig. 5. Energy budget of Workload mice during baseline and workload conditions.
Panel A shows the absolute values and panel B the mass-specific values. To
determine the energy balance we used measures of resting metabolic rate (RMR,
light-grey bars) and daily energy expenditure (DEE). Energy for activity (ACT,
darker grey bars) was calculated by deducting RMR from DEE. In addition, the
gross energy intake (GEI) was calculated on the basis of the absolute food
intake during the DLW measurements. Metabolisable energy intake (MEI, striped
bars) was then calculated from GEI, assuming that digestive efficiency
together with energy lost in the urine was 79.1%
(Hambly and Speakman, 2005 ).
The white bars represent the energetic value of the food that is not
metabolized (GEIMEI=Waste, W). The numbers in the bars represent the
amount of energy (either in kJ d1 or in kJ
g1 d1) spent on each part of the energy
budget. The bracket shows the surplus energy available to the animals for
growth.
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© The Company of Biologists Ltd 2007