First published online November 28, 2008
Journal of Experimental Biology 211, 3871-3878 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.023101
Timing of the daily temperature cycle affects the critical arousal temperature and energy expenditure of lesser long-eared bats
Christopher Turbill*,
,
Gerhard Körtner and
Fritz Geiser
Centre for Behavioural and Physiological Ecology, Zoology, University of
New England, Armidale, New South Wales, 2351 Australia

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Fig. 1. Representative examples of skin temperature (Tskin;
filled circles and dotted line) and metabolic rate (MR; solid line) of captive
bats exposed to a diurnal fluctuation in ambient temperature
(Ta; dashed line) with heating commencing (shown by arrow)
at (A) 06:00 h, (B) 09:00 h or (C) 12:00 h. The photoperiod spanned from 06:00
h to 18:15 h.
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Fig. 2. Histograms showing times of arousal (filled vertical bars) and times of
re-entry into torpor (open vertical bars) by bats over their rest phase during
exposure to a diurnal fluctuation in ambient temperature
(Ta; shown at the top) that commenced heating at (A) 06:00
h, (B) 09:00 h or (C) 12:00 h. Bats were placed into respirometry chambers in
the evening (Ta: 13°C) and all had entered torpor
during the night or around the time of lights on. All bats aroused in response
to passive rewarming when heating commenced at 09:00 h or 12:00 h. When
heating commenced at 06:00 h, bats (n=3) remained in torpor
throughout on 4 of 15 (27%) days. The horizontal black and white bars above
indicate the photophase.
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Fig. 3. Ambient temperature (Ta) at the time of active arousal
by bats receiving passive rewarming from exposure to a daily fluctuation in
Ta when heating commenced at 06:00 h, 09:00 h or 12:00 h
(individuals: filled circles and connecting lines; averages: open circles).
Bats aroused at lower Ta and therefore after less passive
rewarming when heating had commenced later in the day (repeated ANOVA:
F2,22=14.4, P<0.001; Tukey's test: all
P<0.05).
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Fig. 5. Energetic costs of active arousal from torpor as a function of ambient
temperature (Ta) for arousals from minimum
Ta without passive heating (triangles) and for arousals
after diurnal passive heating (filled circles). Passive heating provided a
linear reduction in the average cost of active arousal [solid line: energy
expenditure for active arousal
(kJ)=0.84–0.026xTa (°C);
r2=0.87, P<0.001]. During torpor, passive
heating resulted in a net increase in energy expenditure, which can be
included as a component of the total energy expenditure for arousal [open
circles; dashed line: total energy expenditure for arousal
(kJ)=0.78–0.02Ta (°C);
r2=0.82, P<0.001].
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Fig. 6. Energy consumption over the rest phase (12 h) of bats exposed to a diurnal
fluctuation in Ta when heating commenced at 06:00 h (open
circles), 09:00 h (triangles) or 12:00 h (filled circles) as a function of
time spent normothermic (0 h represents bats that remained in torpor). The
time of commencement of heating had no significant effect on the slope of the
relationship between energy consumption and time normothermic [general linear
modelling (GLM) slope: F2,41=2.4, P=0.10],
however, energy expenditure was greater overall on days when heating commenced
at 06:00 h or 09:00 h, in comparison to 12:00 h [GLM, y-intercept:
F1,43=287, P<0.001; fitted line for combined
06:00 h and 09:00 h: rest phase energy expenditure
(kJ)=0.734.0+0.513xtime normothermic (h); for 12:00 h: energy
expenditure (kJ)=0.261+0.513xtime normothermic (h);
r2=0.90].
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Fig. 7. An approximate model of the change in the critical threshold
Ta that triggers arousal (thermal cue for arousal; thick
dashed line) over the rest phase (daytime) in male N. geoffroyi.
Torpor is entered before or near dawn. Arousal and active rewarming of body
temperature (solid line) from torpor is predicted to occur if and when
Ta (thin dotted line) reaches the thermal cue for arousal.
Bats re-enter torpor in response to cooling of Ta. Torpid
bats arouse near dusk.
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© The Company of Biologists Ltd 2008