First published online October 21, 2005
Journal of Experimental Biology 208, 4099-4108 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01855
Metabolic adjustments to increasing foraging costs of starlings in a closed economy
Popko Wiersma*,
H. Martijn Salomons and
Simon Verhulst
Zoological Laboratory, University of Groningen, PO Box 14, 9750 AA
Haren, The Netherlands

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Fig. 1. Daily energy expenditure in relation to food availability manipulations.
(A) Caloric restriction results in a decrease in energy expenditure. (B)
Increasing foraging costs per reward: when metabolic rate during foraging is
higher than when not foraging, and everything else remains constant (mass,
nocturnal energy expenditure, etc.), daily energy expenditure is an
accelerating function of foraging costs per reward.
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Fig. 2. The hourly averaged metabolic rates (A) and mass-specific metabolic rates
(B) for different food availabilities (night from 0 to 10 h, i.e. 14:00 h to
00:00 h local time). Values are means ±
S.E.M., N=7. The lines are fitted
using 3rd-order polynomials.
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Fig. 3. Body mass in different foraging environments, shown separately for birds
that started in the rich environment (closed symbols, broken lines) and poor
environment (open symbols, solid lines). Arrows indicate treatment order.
Treatment and time had an effect on mass, while order of treatment had not
(Table 1).
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Fig. 4. Association between metabolic rates (measured at ±20.8°C) of
starlings measured during the day (MRday) and the night
(MRnight) during a 24 h respirometer measurement. MRday
and MRnight were strongly correlated (r=0.75,
N=24, P<0.001, controlling for mass); the regression line
is shown. The grey lines show the average values of MRnight for our
birds and the estimated values of MRday in the rich and poor
environment.
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Fig. 6. Energy budgets and flight times in different environments. Also shown (far
right) is the hypothetical budget and flight time in the poor environment of
starlings that maintain high body mass the same as in the rich environment.
Eflight is the energy spent on flying, BMRnight
is the energy spent during the night on BMR only, Enight
is the total energy spent during the night, and Enonfly
day is what is spent during day-time when not flying. The broken lines
in the hypothetical energy budget indicate the surplus energy that would have
been spent due to the extra time spent flying (and less time spent not
flying), and the extra energy that is spent on flying due to the body mass
increase. The total daily energy expenditure (DEE) budget is shown on top of
each bar.
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© The Company of Biologists Ltd 2005