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Journal of Thermal Biology 37 (2012) 392–396

Contents lists available at SciVerse ScienceDirect

Journal of Thermal Biology

journal homepage: www.elsevier.com/locate/jtherbio

Ambient temperature-dependent thermoregulatory role of REM sleep

Deependra Kumar 1, Velayudhan Mohan Kumar 2, Hruda Nanda Mallick n,1
Department of Physiology, All India Institute of Medical Sciences, Ansari Nagar, New Delhi 110029, India

a r t i c l e i n f o a b s t r a c t

Article history: Changes in ambient temperature produce complex effects on sleep–wakefulness. In order to find out
Received 31 October 2011 the mechanisms involved in temperature-sensitive changes in sleep in rats, their thermal preference,
Accepted 14 February 2012 body temperature and sleep were studied before and after the destruction of both peripheral and
Available online 3 March 2012
central warm receptors, by systemic administration of 375 mg/kg capsaicin. Though the pre-treated
Keywords: rats preferred to stay mostly at the ambient temperature of 27 1C, post-treated rats strayed freely into
Capsaicin chambers having ambient temperature of 30 1C and 33 1C. Sleep and body temperature of these rats
Sleep were studied for six hours each, when they were kept at an ambient temperature of 18–36 1C. Total
Body temperature sleep time, especially REM sleep, was maximum at 30 1C in pre-treated rats, but this REM sleep peak at
Ambient temperature
30 1C disappeared after capsaicin administration. Body temperature increased sharply in post-treated
Thermal preference
rats, at ambient temperatures above 30 1C. Apart from the ability to defend body temperature at high
ambient temperature, avoidance of warm ambient temperature and increase in REM sleep are the
behavioral measures which are lost in post-treated rats. Results of this study suggest that the ambient
temperature-related increase in REM sleep at 30 1C could be part of the thermoregulatory measures.
& 2012 Elsevier Ltd. All rights reserved.

1. Introduction receptors. Moreover, those reports are based on the results

Ambient temperature (Tamb) does influence sleep in animals,
including rats. It is generally assumed that one sleeps more at a
comfortable temperature. When rats are given the choice of
different Tamb to select from, they prefer to stay in 27 1C Tamb
(Gulia et al., 2005; Kumar et al., 2009; Ray et al., 2004, 2005).
However, they had maximum total sleep time (TST), when they
were kept at 30 1C (Gulia et al., 2005; Kumar et al., 2009;
Mahapatra et al., 2005; Sakaguchi et al., 1979; Szymusiak and
Satinoff, 1981). This influence of warm Tamb on sleep is probably
mediated by warm receptors, as the increased Tamb does not
produce a significant change in sleep of rats with destroyed warm
receptors (Benedek et al., 1980; Obal et al., 1983). The tempera-
tures preferred by the rats before and after the destruction of
warm receptors were not studied in these reports. It is possible
that the change in thermal preference of the rats was responsible
for the lack of sleep changes in rats with destroyed warm
Abbreviations: Ambient temperature, Tamb; Body temperature, Tb;
Sleep–wakefulness, S–W; Total sleep time, TST; Slow wave sleep, SWS
n
Corresponding author. Tel.: þ91 11 26594881;
fax: þ 91 11 26588641, þ91 11 26588663.
E-mail addresses: kdeepen@gmail.com (D. Kumar),
wfsrs2005@rediffmail.com (V.M. Kumar), drhmallick@yahoo.com (H.N. Mallick).
1
Tel.: þ91 11 26594623.
2
Present address: Comprehensive Centre for Sleep Disorders, Sree Chitra
Tirunal Institute for Medical Sciences and Technology, Thiruvananthapuram
695012, India. Tel.: þ 91 471 2328004; fax: þ 91 471 2341814.
obtained from two separate groups of rats (normal vs. capsai-
cin-treated) and the animals were exposed only to two Tamb. It is
also necessary to test the changes in different components of
sleep before and after the destruction of warm receptors. The
interrelationship between the changes in sleep and body tem-
perature (Tb) is yet another area which has not been studied in
capsaicin-treated animals.
Capsaicin, an active substance of hot peppers, can stimulate
the warm receptors, when given in small doses, and can selec-
tively destroy the warm receptors, when given in large doses. So,
it is widely used in studies on thermoregulation (Szolcsányi and
Jancsó-Gábor, 1973). It produces a steep fall in body temperature
(Tb) when small quantity (test dose) is subcutaneously injected in
rats (Jancsó-Gábor et al., 1970a). However, after the warm
receptors are destroyed with large systemic doses of capsaicin,
no hypothermic action is produced on administration of a test
dose of this drug (Jancsó-Gábor et al., 1970a, 1970b). In the
present study, a small quantity of capsaicin was subcutaneously
injected in rats to find out the functional integrity of warm
receptors, and large doses were administered to destroy all the
warm receptors. The thermal preferences of the animals were
studied before and after warm receptors were destroyed. Capsai-
cin sensitivity test was also performed before and after large
doses of capsaicin injection. TST, REM sleep, slow wave sleep
(SWS) and Tb at wide ranging Tamb were analyzed in order to
understand the interrelationship between Tamb, sleep, and Tb in
these animals.

0306-4565/$ - see front matter & 2012 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jtherbio.2012.02.005
 Author's personal copy
D. Kumar et al. / Journal of Thermal Biology 37 (2012) 392–396
2. Materials and methods
Ten adult male Wistar rats, weighing between 250 and 275 g,
were chronically implanted with electrodes for electroencephalo-
gram (EEG), electromyogram (EMG) and electrooculogram (EOG),
as described elsewhere (Kumar et al., 1984). A Radio-transmitter,
TA10TA-F40 (Data Science International, USA), was implanted in
the abdomen of the animals to record Tb (Vetrivelan et al., 2003).
All procedures were conducted in accordance with the rules of the
Committee for the Purpose of Control and Supervision of Experi-
ments on Animals (CPCSEA) and were approved by the Institu-
tional Animal Ethics Committee, AIIMS, New Delhi, India. The rats
were maintained at 14 h light and 10 h dark conditions with light
on from 06:00 h, with controlled temperature (2571 1C), and ad
libitum access to food and water.
The recording of various parameters on different days is shown
in Fig. 1. After a ten-day recovery period and two-day habituation
in the recording chamber, the functional integrity of the warm
receptors in these rats was assessed using the capsaicin sensitiv-
ity test (Jancsó-Gábor et al., 1970a, 1970b). This was done by
injecting capsaicin (8-Methyl-N-Vanillyl-6-Nonenamide, Sigma,
USA) subcutaneously at a dose of 2 mg/kg body weight and
measuring the fall in Tb.
The thermal preference of the rats was studied for six hours
(11:00–17:00 h) in an environmental chamber with three com-
partments, fitted with activity monitoring systems (Habitest
response sensing module, model E45-04, Coulbourn Instrument,
Fig. 1. Schematic diagram shows experimental steps conducted on different days.
393
USA). Vibration-sensing platforms were kept below the floors of all
the compartments of the environmental chamber (Fig. 2, upper
panel). The vibration produced by the movement of the rat was
monitored using E24-72 resistive bridge transducer monitor. The
sensitivity and threshold of the transducer were adjusted with the
animals’ weight. All transducers were connected to the interface
card in the computer (controlling the three vibration sensing
platforms) and finally to the Graphic state program. Each move-
ment of the animal was registered as a count (from that particular
platform/compartment) in Graphic state software. Thus, the soft-
ware assessed the time interval between first count from one
compartment and first count in other compartment i.e. time spent
by the animals in each of the compartments. The three compart-
ments of the chamber were maintained at 271, 301, and 33 1C
respectively. These three Tamb were selected on the basis of the
existing literature; 27 1C being the Tamb preferred by the rats as per
the earlier report (Gulia et al., 2005; Kumar et al., 2009; Ray et al.,
2004, 2005) and 30 1C and 33 1C being the temperatures shown to
induce maximum sleep (Gulia et al., 2005; Kumar et al., 2009).
Sleep–wakefulness (S–W) and Tb were recorded for six hours
(11:00–17:00 h) in each of the rats at 181, 211, 241, 271, 301, 331
and 36 1C Tamb, on alternate days. Flexible cables with connectors
were plugged to the rats’ heads (Fig. 2, lower panel). Their outputs
were connected to a digital recorder (BSL PRO 36, BIOPAC system
Inc., USA) for recordings EOG, EEG, and EMG, along with tele-
metrically assed Tb (DATA QUEST 1.1, Data Science International,
and USA). The conditions of the recording cage were kept
identical to that of the thermal preference recording chamber,
except for Tamb, which varied on different days of exposure. The
temperature regulating system of the recording cage was capable
of obtaining the altered Tamb within five minutes. The relative
humidity varied from 70% to 80%.
Capsaicin was systemically administered to destroy the warm
receptors throughout the body. Total dose of 375 mg/kg capsaicin
was given subcutaneously over a period of four days, under light
ether anesthesia. Two injections each were given at intervals of
20–30 min, on all the four days. On the first day, the rats were given
two injections of capsaicin at the dose of 25 mg/kg. They received
25 mg/kg and 50 mg/kg, on the second day. Two injections of
50 mg/kg were given on the third day, and on the fourth day
50 mg/kg and 100 mg/kg were administered. The first injection of
capsaicin on each day was preceded by an intraperitoneal injection
of atropine (25 mg/kg). Results mentioned in this study are based on
the findings from five rats that survived the injection procedures.
Three weeks after the last capsaicin injection, the destruction
of warm receptors was assessed (functionally) on the basis of Tb
response to capsaicin test dose. The thermal preferences, S–W
and Tb of the capsaicin-treated animals were studied at Tamb
similar to those of the pre-treated animals. At the end of the
study, capsaicin sensitivity was reassessed in the treated animals.
The S–W records were split into 15 s epochs and visually
classified on the basis of EEG, EMG and EOG, as described earlier
(John et al., 1994). The Tb and percentage TST, REM sleep and SWS
(calculated as percentage of total recording time) were compared
by non-parametric two-way analysis of variance (Friedman’s
test). The S–W and Tb data at 27 1C were taken as the control
reading for comparison (using Wilcoxon Signed Rank Test) with
the data of these parameters obtained at 181, 211, 241, 301, 331
and 36 1C. Data from two consecutive Tamb, and before and after
destruction, were also compared using the same test.
3. Results
The subcutaneous injection of 2 mg/kg of capsaicin (capsaicin
sensitivity test) in normal rats produced a fall of  3 1C in Tb and
 Author's personal copy

394 D. Kumar et al. / Journal of Thermal Biology 37 (2012) 392–396

Fig. 2. Schematic diagram shows recording setup used for assessing thermal preference (upper panel), sleep–wakefulness and body temperature (lower panel).

Fig. 3. The figure shows the mean 7 S.D. of body temperature of pre-treated and
post-treated rats (n¼5) when 2 mg/kg subcutaneous capsaicin was injected.
n
po 0.05, significant change after test dose injection compare with before Tb in
pre-treated rats. zp o0.01, significant change compared between pre and post-
treatment of capsaicin.
Fig. 4. The figure shows thermal preference of rats (n¼5) recorded for 6 h (11:00–
17:00 h). Y-axis shows the duration of stay (mean 7 S.D.) in each compartment,
pre and post-injection of capsaicin. X-axis shows temperatures in the three
compartments. npo 0.05, significant change between thermal preference at
27 1C and 30 1C in pre-treated rats. np o 0.05, significant change between thermal
preference at 27 1C and 33 1C in pre-treated rats. yp o0.05, significant change
between pre- and post-injection of capsaicin.

The pre-treated rats preferred to stay at 27 1C, (compared to

the hypothermia persisted for about an hour (Fig. 3). Three weeks
after the destruction of warm receptors, with systemic injection
of high dose of capsaicin, subcutaneous injection of 2 mg/kg of
capsaicin did not produce any fall in Tb. These rats remained non-
responsive to capsaicin test till the end of the study.
30 1C, z ¼2.023, p¼0.043, and 33 1C, z¼2.023, p ¼0.043, two
sided) during 11:00–17:00 h, when they were provided a choice
of three different temperatures of 271, 301, and 33 1C (Fig. 4). After
a high dose injection of capsaicin, there was a significant decrease
(z ¼2.023, p ¼0.043, two sided) in the preference for 27 1C, and an
 Author's personal copy

D. Kumar et al. / Journal of Thermal Biology 37 (2012) 392–396 395

Fig. 5. The figure shows the mean 7 S.D. of REM sleep, slow wave sleep and total sleep time (bar diagram) and body temperature (line diagram) of rats (n¼ 5) pre- and
post-treatment of capsaicin when they are exposed to various ambient temperatures. X-axis shows the ambient temperatures. Y-axis shows percentage recording time (bar
diagram) and body temperature (line diagram). np o 0.05, significant change compared to 27 1C. yp o0.05, zp o 0.01, significant change between pre- and post-treatment of
capsaicin. np o 0.05, significant change in pre-treated rats when compared between two consecutive ambient temperatures. mpo 0.05, significant change in post-treated
rats when compared between two consecutive ambient temperatures.

increase (z ¼2.023, p¼0.043) in the preference for 30 1C. But the
increase in preference of 33 1C was not significant (z ¼1.753,
p ¼0.080).
In the pre-treated rats, the average TST at 27 1C Tamb was
67.91 77.8% of 11:00–17:00 h recording time (Fig. 5). TST was the
maximum at 30 1C, (69.89 710.8% z¼ 2.023, p¼0.043, two sided);
it decreased above (at 36 1C, 49.71 76.27%, z ¼2.023, p¼ 0.043)
and below (at 18 1C, z ¼2.023, p ¼0.043 and 21 1C, z ¼2.023,
p ¼0.043) this Tamb. There were no significant differences in TST
at 24 1C (z ¼0.674, p ¼0.500) and 33 1C (z¼0.674, p¼0.500) when
compared with 27 1C. The TST peak at 30 1C was primarily due to
the increase (16.0873.1%, z¼2.023, p¼ 0.043) in REM sleep
(Fig. 5). The REM sleep peak at 30 1C disappeared (10.2672.4%,
z¼ 0.944, p¼0.345) in post-treated rats. In addition, the TST peak
was observed at 33 1C (69.43 78.1%, z¼2.023, p ¼0.043) in these
rats. Though the mean Tb increased significantly (37.7670.3 1C,
z¼ 2.023, p ¼0.043) at 36 1C in pre-treated rats, there was a steep
rise in Tb above 30 1C (at 33 1C, Tb ¼38.14 70.6 1C, z ¼2.023,
p ¼0.043 and 36 1C, Tb ¼38.93 70.4 1C, z¼ 2.023, p ¼0.043) after
treatment (Fig. 5).
known for a long time that systemic injection of high dose of
capsaicin in the animals produced insensitivity to subsequent test
dose injections for months (Jancsó-Gábor et al., 1970a; Szolcsányi
and Jancsó-Gábor, 1973). Moreover, electrophysiological and
morphological studies have shown that high dose of subcuta-
neous capsaicin administration produces long-lasting impairment
of peripheral and central warm receptors (Hori, 1981; Hori and
Tsuzuki, 1981; Jancsó-Gábor et al., 1970a, 1970b; Obál et al.,
1979, 1980; Szolcsányi, 1977; Szolcsányi et al., 1971; Szolcsányi
and Jancsó-Gábor, 1973). In the present study also, these animals
were unable to defend Tb, as evident from the sharp increase in Tb
at Tamb above 30 1C. Similar findings were reported earlier where
rats succumbed to heat stroke at Tamb that the normal rats could
easily cope with (Benedek et al., 1980; Obál et al., 1979).
Capsaicin-treated animals also showed some fall in Tb in cold
Tamb as reported in an earlier study (Benedek et al., 1983). Thus, a
high dose of capsaicin administration that destroyed warm
receptors, grossly reduced the Tb regulation at higher Tamb, though
thermoregulation in lower Tamb was also affected.
4.2. Effect of capsaicin treatment on thermal preference

4. Discussion
4.1. Effect of capsaicin treatment on Tb regulation
It can be safely assumed that the warm receptors are sub-
stantially destroyed in the rats treated with high dose of capsai-
cin, as there was no fall in Tb to capsaicin sensitivity test. It was
Control rats did not venture into Tamb above 27 1C, but those
with destroyed warm receptors moved freely into 30 1C and 33 1C
Tamb. Preference for Tamb above 27 1C after capsaicin treatment
can be considered as an impairment of thermal selection behavior
of the animals (Obál et al., 1981). Earlier studies have shown that
the thermal selection by the animals is preserved after selective
destruction of peripheral warm receptors (Gulia et al., 2005).
 Author's personal copy
396 D. Kumar et al. / Journal of Thermal Biology 37 (2012) 392–396
In the light of this earlier report, it can be assumed that the
central warm receptors are primarily responsible for the changed
thermal selection behavior of the rats in the present study.
4.3. Effect of capsaicin treatment on S–W regulation
Several studies suggested that S–W in rats is sensitive to Tamb
(Gulia et al., 2005; Kumar et al., 2009; Schmidek et al., 1972;
Szymusiak and Satinoff, 1981). A moderately warmer Tamb (30 1C)
promotes sleep in normal rats (Gulia et al., 2005; Kumar et al.,
2009; Sakaguchi et al., 1979; Szymusiak and Satinoff, 1981). This
increase in sleep is probably a thermoregulatory response to
thermal load (Obal et al., 1983).
It is interesting to note that TST increased relatively
(69.4378.1%, p¼0.043) at 33 1C Tamb, along with a steep increase
in Tb, after destruction of warm receptors. In the absence of
peripheral and central warm receptors, induction of sleep at 33 1C
should be attributed to a direct effect of increase in Tb, and brain
temperature. Lower Tamb did not significantly change sleep,
despite the small reduction in Tb. Thus it is sleep at higher Tamb
that is mainly affected in capsaicin-treated rats.
The effect of Tamb on REM sleep was more marked, and it was
maximum at 30 1C in control rats. REM sleep has been considered
as a state where thermoregulation ceases (Permeggiani et al.,
1977). But it is possible that Tamb-related increase in REM sleep in
control rats, specifically at 30 1C, could be part of thermoregula-
tory measure. Decreased muscle tone during REM sleep leads to
decreased heat production by the muscles, which may help in
decreasing the thermal load (Aristakesian and Vataev, 1993). The
selective increase in REM sleep (REM sleep peak) at 30 1C
disappears after the destruction of the warm receptors. This
indicates that the animal lost REM sleep-related heat dissipating
mechanism at warm Tamb. Our finding is in agreement with
various studies on capsaicin-treated rats in which heat defense
responses such as skin vasodilation (Dib, 1983; Jancsó-Gábor
et al., 1970a, Obál et al., 1982), salivation (Jancsó-Gábor et al.,
1970a; Obál et al., 1979), prone body extension, heat escape
locomotion (Obál et al., 1979), skin cooling operant behavior (Dib,
1983; Hori and Tsuzuki, 1981) and water bathing (Obál et al.,
1980) are lost after capsaicin treatment.
5. Conclusion
Sleep, especially REM sleep, was increased at 30 1C in the
normal rats, though they preferred 27 1C Tamb. It can be assumed
from the present study that thermal selection behavior and
increase in REM sleep at 30 1C Tamb are dependent on central
warm receptors. Both these responses are used for thermoregula-
tion, but they are not interrelated physiological responses.
Increase in REM sleep at 30 1C Tamb in normal rats could be
considered as a thermoregulatory function of warm receptors,
which does not involve conscious perception of warm sensation.
Acknowledgment
This study was supported by the Defense Research and Devel-
opment Organization of India.
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