Physiology & Behavior. Vol. SY. No.

2, pp 277-2X1, 15~96
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0031-9384(95)02127-2

Effects of Short- and Long-Term REM Sleep

Deprivation on Sexual Behavior in Male Rats
JAVIER VELAZQUEZ-MOCTEZUMA,
EMIL10 DOMINGUEZ
AND SOCORRO RETANA-MARQUEZ
Departamento

SALAZAR

de Biologia de la ReproduccGn, Universidad Aut&oma Metropolitana-Iztapalapa,

Col. Vicentina, C.P. 09340, Del. Iztapalapa, Mkxico City, D.F., M&co
Received 23 February

VELAZQUEZ-MOCTEZUMA

J., E. DOMINGUEZ

Purisima y Michoacan,

1995

SALAZAR

AND S. RETANA-MARQUEZ.
Eflects ofshoutBEHAV 59(2) 277-281,
1996.-The
influence of selective REM sleep deprivation on masculine sexual behavior has been a matter of
controversy. In the present study, the sexual behavior of male rats was analyzed in subjects deprived of REM sleep by
the island technique for 24 or 16 h daily during 20 days. When compared to control rats, both groups displayed
changes in sexual performance
since the first day. The effects were: an increase in mount, intromission and
ejaculation latencies and in mount frequency as well; a decrease of ejaculation frequency and of the Hit rate. The
effects became stronger as REM sleep deprivation progressed. Rats deprived of REM sleep for 24 h were extremely
debilitated after 12 days and some of them died, whereas the rats REM deprived for 16 h remained healthy during the
20 days. These data indicate that REM sleep deprivation interferes with the mechanisms that regulate male sexual
behavior.

and long-term REM sleep deprivation on sexual behavior in male rats. PHYSIOL

REM sleep

REM sleep deprivation

Sexual behavior

Masculine sexual behavior

tion in animals has been the island technique (8). Nevertheless, as

this technique implies submitting the animals to a stressful
situation, a reliable control group for the stress component is
needed. Most of the authors have used a large platform control
group in which the rats can reach the REM sleep stage (24). A
wet control group in which the animals are submitted to immersion in cold water, has also been used (21). Both procedures,
however, have been part of the controversy because the results
have often been different from the unmanipulated control.
In the present study we analyzed the effect of short- and long
term REM sleep deprivation on sexual performance in male rats.
In addition, we examined a new attempt to assess the stress
component in REM deprivation experiments,
with a control
group which was REM deprived in exactly the same condition,
but only for 16 h a day, allowing them to sleep for the remaining
8 h.

INTRODUCTION

THE SELECTIVE deprivation of the Rapid Eye Movement

(REM) sleep stage has been used for several years in an attempt
to elucidate the function of this stage in the organism (for review
see: 25). To date, it is clear that REM sleep deprivation can
induce alterations in some behavioral patterns and mainly in
those that are under hormonal control, like aggression (6), fear
(51, drinking (9) and eating (9,12). Regarding sexual behavior,
however, some controversial papers have been reported. Morden
et al. (14), described that low copulators
increased their
performance when they were REM deprived. Our group has
reported that the effect of steroid hormones on sexual behavior is
modified by REM deprivation
both in females and males
(3,22,23). On the other hand, Hicks et al. (71, reported that 4 days
of REM deprivation did not induce significant changes of masculine sexual performance
Another source of controversy in REM deprivation studies
refers to the stress component present in all the techniques used
for selective REM sleep deprivation (4,131. Concerning sexual
behavior, it has been reported that acute stressful situations have
a facilitatory effect on rat male sexual performance (1,ll). In
addition, the effects of stress on sexual behavior seem to depend
on the nature and the characteristics of the stressor (17).
To date, the most common technique used for REM depriva-

METHODS

Adult male Wistar rats, bred in our vivarium, were used in

this study. Animals were tested twice for masculine sexual
behavior and the active rats were randomly assigned to one of the
following groups:
Group A: N = 10. REM deprived (REMd) group. The animals
were submitted to the island technique which, in brief, consists in

To whom requests for reprints should be addressed.

277

278

VELAZQUEZ-MOCTEZUMA,

ICONTROL

SALAZAR AND RETANA-MARQUEZ

placing the animal on a circular platform (4.5 cm diameter.)

surrounded by water up to 1 cm below the surface of the
platform, with ad lib access to food and water. This platform was
located in the middle of a standard acrylic cage. The height of the
platform was decreased from 10 cm, of the original technique, to
4.5 cm, to avoid the immobilization component of the original
technique. This lower water level (3.5 cm), allowed the animal to
come down and walk around the cage. The water was changed
daily. Animals remained in the cage for 20 days and were taken
out only for sexual behavior studies.
Group B: N = 10. Partially REM deprived (PREM) group. The
subjects in this group were submitted to the island technique but
only for 16 h a day. The remaining 8 h, the animals were allowed
to sleep in a dry cage. As rats normally concentrate their sleep in
the light period, these control rats were allowed to sleep during
the light period.
Group C.N = 10. Control group. These animals remained in the
vivarium under standard laboratory conditions during the entire
experiment.
All the rats were maintained in a room with controlled
temperature and ventilation, and under a 12/12 inverted light
cycle (lights off: 09 h. Lights on: 21 h.). Food and water were
available Ad Lib.
Male sexual behavior tests were done during the dark phase of
the cycle on days 1, 4, 8, 12, 1.5, and 20. In brief, males were
placed in a circular arena (45 cm diameter) for a 5 min habituation period, then a receptive female rat was presented. The
stimulus female was ovariectomized and 2 wk later, was treated
with estradiol benzoate (5 pg/.l
ml corn oil) followed by
progesterone (2 mg/.l
ml), administered 48 and 4 h, respectively, before the behavioral test.
During the sexual behavior test, the following parameters
were recorded (for review see: 10 and 18):
Mount Latency (ML): the time that elapsed between the presentation of the female and the first mount without intromission.
Intromission Latency (IL): The time that elapsed between the
presentation of the female and the first mount with intromission.
Ejaculation Latency (EL): The time that elapsed between the first
intromission and ejaculation.
Postejaculatory
Period (PR): The time that elapsed between
ejaculation and the first mount of the next copulatory series.
Mount Number (MN): Number of mounts preceding ejaculation.
Intromission Number (IN): Number of intromissions preceding
ejaculation.
Ejaculation Frequency (EF): Number of ejaculations displayed in
30 min.
Hit Rate (HR): number of mounts plus number of intromissions
divided by the number of intromissions.
Average Interintromission
Interval (III): Ejaculation Latency divided by the number of intromissions.
Average Intercopulatory Interval (ICI): Ejaculation Latency divided by the number of intromissions plus the number of mounts.
Statistical analysis was performed by means of the KruskalWallis one way ANOVA followed by Mann-Whitney
U-tests.

12

DAYS
MREMd16

DOMINGUEZ

mREMd24

FIG. 1. Mount (A), Intromission (B), and Ejaculation (C) Latencies of

male rats submitted to partial (16 h/day) or total (24 h/day) REM sleep
deprivation during 20 consecutive days. Tests were done only on the days

indicated.
Mann-Whitney U-test compared to control.
* = p < 0.05.
** = p < 0.01.
*** = p < 0.005.

TABLE 1
PERCENTAGE OF RATS PRESENTING MOUNTS CM), INTROMISSIONS (I) AND EJACULATIONS (E).
x INDICATES PERCENTAGE OF DEAD ANIMALS
COlltrOl
Day
1

4
8
12
15
20

REMd24

REMd16

100
100
100
100
100
100

100
100
100
100
100
100

100
100
100
100
100
100

0
0
0
0
0
0

100
100
100
100
100
100

100
100
100
100
100
100

100
100
100
100
90
100

0
0
0
0
0
0

90
100
100
90
20
_

90
100
100
70
10
_

90
100
90
50
0

0
0
0
10
30
50

REM DEPRIVATION

AND SEXUAL

BEHAVIOR

279

RESULTS

TABLE 2

As can be seen in Table 1, since day 12 the REMd group

showed significantly lower percentages of subjects presenting
mounts, intromissions and ejaculation when compared either with
the PREM group or with the control group as well. It must be
noted that some experimental subjects died during the procedure
and the remaining subjects showed a marked deterioration of
their health. However, on day 20 the behavioral test could not be
done mainly due to the aggressive behavior displayed by the
male towards the female. On the other hand, subjects under
partial REM sleep deprivation showed no fatalities and all of
them remained sexually active during the 20 days of the experimental procedure.
The results obtained regarding mount, intromission and ejaculation latencies are shown in Fig. 1. The REMd group showed a
significant increase in both mount and intromission latencies
during the first day (LM, Kruskal-Wallis;
H = 8.5 p < 0.01. LI,
Kruskal-Wallis;
H = 7.86, p < 0.02). An important increase in
intromission
latency was also observed during day 12. The
PREM group showed only a significant decrease in mount latency on the first day and an increase in intromission latency on
day 12 as well. Ejaculation latencies showed a marked increase
on day 12 (Kruskal-Wallis H = 10.83, p < 0.001) in both PREM
and REMd groups. As has been mentioned, the experimental
group showed no ejaculations during day 15. The PREM group
also displayed a significant increase of ejaculatory latency on day
20 (Kruskal-Wallis H = 12.37, p < 0.001).
Concerning the number of mounts preceding ejaculation both
the REMd and the PREM group showed significant increases in
almost all of the recording days (Kruskal-Wallis H = 12.45 p <
0.001) (Fig. 2). The number of intromissions preceding ejacula-

DATA WERE OBTAINED FROM THE FIRST

EJACULATORY SERIES AND ARE EXPRESSED AS MEAN
(S) + SEM MANN-WHITNEY
U-TEST COMPARED
TO CONTROL GROUP
Average Intercopulatory Interval
Day
1

4
8
12
15
20

C0ntr0l

38.5
30.38
31.1
26.3
35.9
36.0

& 2.58
* 4.0
+ 2.9
of-4.0
f 3.4
c 34

47.3
36.1
39.8
31.2
44.2
44.1

* 4.15
i_ 4.5
_+4.6
* 4.8
i 5.1
+ 5.2

REMd 16
34.6

4.4

30.5
26.8
41.0
37.9
3x.7

&
f
+
f
*

7.6
1.7
l.h*
5.6
2x

54.4
54.2
54.6
X2.6
77.2
86.1

&
i_
&
+
+
i_

5.6
h s*
5.7
Y.l$
I I x*
7 4$

Day

1
4
8
12
15
20

REMd24

24.94
31.72
20.35
72.2

*
+
i
*

2 3w
2.50
2.30t
2.90

REMd24

64.45
49.5
57.x
61.X4

f 7.18
+ 5.9
t Il.)*
+ 12.4*

* = p < 0.05; : = p < 0.01; $ = p c I) OOI

tion, however, did not show significant differences among all the
groups.
On the other hand, ejaculatory frequency showed a significant
decrease in the REMd group from the first day of the manipulation throughout the entire experiment (Kruskal-Wallis H = 13.76
p < 0.001). The PREM group showed a trend to decrease the
frequency from the first day also, but it reached significant values
starting on day 12 (Fig. 3A). The postejaculatory
period was

DAYS
BlREMdl6

-REMd

of mounts preceding the first ejaculation in male rats submitted

during 20 consecutive days. Tests were done only on the days indicated.
Mann-Whitney
U-test compared to control.
* = p < 0.05.
** =p < 0.005.
*** =p < 0.001.

to partial (16 h/day)

OCONTROL
FIG. 2. Mean number

24
or total (24 h/day)

REM sleep deprivation

VELAZQUEZ-MOCTEZUMA,

280

TABLE 3

Hit Rate

4
8
12
15
20

REMd16

COlltrOl

0.87
0.80
0.79
0.84
0.80
0.85

f
f
*
f
k
k

0.03
0.02
0.10
0.04
0.02
0.03

0.64
0.59
0.53
0.50
0.52
0.46

f
f
f
f
f
f

0.03t
0.002t
0.002*
0.001t
0.003*
O.OOlt

REMd24

0.40
0.69
0.44
0.39

SALAZAR AND RETANA-MARQUEZ

Table 2 shows the results obtained regarding the intercopulatory and the interintromission
intervals. As can be seen, the
PREM group showed a marked increase in the interintromission
interval during the last part of the experiment (days 12, 15, 20;
Kruskal-Wallis
H = 13.2, p < 0.001). The Hit Rate (Table 3)
was affected in both REMd and PREM group with significant
decrease from the first day until the last day (Kruskal-Wallis
H = 7.5 p < 0.02).

DATA WERE OBTAINED FROM THE FIRST

EJACULATORY SERIES. MANN-WHITNEY
U-TEST
COMPARED TO CONTROL GROUP

Day

DOMINGUEZ

+ 0.09t
f 0.003
f 0.06t
k 0.061
_
_

DISCUSSION

The present results strongly suggest that the selective deprivation of the REM sleep stage induces a major deterioration of
masculine sexual behavior in male rats. This effect was worse
with chronically applied REM sleep deprivation (15 days) but
started in the first 24 h. As has been reported elsewhere (16),
REM sleep deprivation by the island technique is an aggressive
procedure that, if prolonged more that 10 days, can cause the

* = p < 0.025: t = p < 0.005.

significantly modified during behavioral tests corresponding

to
days 1, 4 ,8 and 12 in both the REMd and PREM groups
(Kruskal-Wallis H = 8.12 p < 0.01) (Fig. 3B).

DAYS
IZICONTROL

mREMd

16

-REMd

24

FIG. 3. (A) Ejaculatory Frequency during a 30 min test and (Bl Postejaculatory Period after the first ejaculatory series in male rats submitted to partial
(16 h/day) or total (24 h/day) REM sleep deprivation during 20 consecutive days. Tests were done only on the days indicated.
Mann-Whitney u-test compared to control.
* =p < 0.05.
**=p<o.o1.
*** =p < 0.005.

REM DEPRIVATION

AND SEXUAL

281

BEHAVIOR

death of the experimental subjects. This well known debilitating

effect of REM sleep deprivation was observed (16) and some
animals died after more than 10 days of the procedure. No
anatomical cause of death was identified. Nevertheless, the remaining rats did not show any motor impairments that could
prevent the display of the complete pattern of masculine sexual
behavior. These results indicate that REM deprivation by itself
interferes with masculine sexual behavior mechanisms aside from
its debilitating effect.
Laboratory rats normally sleep about 13 h per day and most of
this time is during the light part of the cycle. Between H-20% of
the sleep time corresponds to the REM sleep stage. Daily REM
sleep deprivation limited to 16 h per day was sufficient to induce
changes in sexual behavior from the first day. Thus, these data
demonstrate that, at least for sexual behavior, this procedure is
not suitable as an acceptable control for REM sleep deprivation
studies. On the other hand, the results indicate that even partial
REM sleep deprivation can elicit modifications of sexual behavior similar to those induced by total REM deprivation but,
without its lethal consequences. Thus, it may be possible that, in
the study of some behaviors, partial REM sleep deprivation can
be used instead of total REM sleep deprivation. Moreover, it
seems that partial REM sleep deprivation is a procedure to which
the animal can adapt easier than to total REM sleep deprivation.
In a previous study on masculine sexual behavior, we analyzed the results obtained when rats were submitted to different
stressful situations (17). The modifications of sexual behavior
induced by stress seem to depend on the nature of the stressful
situation. The effects induced by REM deprivation, both total and
partial, have some similarities to the effects obtained by immersion in cold water for 15 min, whereas the effects on sexual
behavior induced by electric foot shocks or by immobilization are

quite different from the effects observed in REM deprived animals.

It has been reported that animal models of depression, in
which pleasure-seeking behaviors are diminished, can be induced
by chronic stress (26) or by an unavoidable stressful situation, the
so called learned helplessness (19). The motivational component
of sexual behavior is mainly reflected by mount latency and the
average intercopulatory interval. In the present study, however,
these parameters were sparsely affected. Thus, it seems that
neither partial nor total REM sleep deprivation are inducing a
syndrome of depression.
Using an innovative technique, Rechtshaffen
et al.. (2,151
have published a series of studies on REM sleep deprivation. ln
their experience, the syndrome displayed by the REM deprived
rat has its own particular features (9) and should not be confounded with the effects induced by stress (16), at least when
confronted with the classical response to stressful situations,
originally reported by Selye (201. Similar results were obtained in
a comparative study using, besides the island technique, multiple
platforms and the pendulum technique. No classical signs of
stress were detected in any of these deprivation procedures. The
authors suggested that REM deprivation is only a mild stressor
(4).
Thus, it seems that REM sleep deprivation affects masculine
sexual behavior in rats, but the extent of the participation of
stress in this effect remains to be elucidated.
ACKNOWLEDGEMENTS

The authors want to express their gratittude to Ms. Edith Monroy

Lopez for the reviewing of the manuscript. This work was partly supported by DGICSA, Grant Number 911573 and by CONACyT, Grant
Number D0245N9201
and 400200-l 703.M9207.

REFERENCES
1. Barfield, R.; Sachs, B. Sexual behavior stimulation by painful electrical shock to skin in male rats. Science 161:392-395;
1968.
7. Bergmann, B. M.; Kushida, C. A.; Everson, C. A.; Gilliland, M. A.;
Obermeyer, W.; Rechtshaffen,
A. Sleep deprivation in the rat: II.
Methodology. Sleep 12:5-12; 1989.
J. REM sleep
3. Canchola, E.; Monroy, E.; Velazquez-Moctezuma
deprivation facilitates the estrogen effects on heterotypical
sexual
behavior in male rats. Physiol. Behav. 37:33-37; 1977..
4. Coenen, A. M. L.; Van Luijtelaar, E. L. J. M. Stress induced by three
procedures
of deprivation
of paradoxical
sleep. Physiol. Behav.
35:501-504;
1985.
5. Hicks, R. A.; Moore, J. D. REM sleep deprivation diminishes fear in
rats. Physiol. Behav. 22:689-692;
1979.
6. Hicks, R. A.; Moore, .I. D.; Hayes, C.; Phillips, N.; Hawkins, J. REM
sleep deprivation
increases aggressiveness
in male rats. Physiol.
Behav. 22:1097-1100;
1979.
7. Hicks, R. A.; Bautista, J.; Phillips, N. REM deprivation does not
increase the sexual behaviors of male rats. Percept. Motor Skills
73:127-130;
lY91.
8. Jouvet, P.; Vimont, P.; Delorme, F.; Jouvet, M. Etude de la privation
selective de la phase paradoxale de sommeil chez le chat. CR Sot.
Biol. (Paris) 158:756-759;
1964.
9. Kushida. C. A.; Bergmann, B. M.; Rechtschaffen, A. Sleep deprivation in the rat: IV. Paradoxical sleep deprivation. Sleep 12 (1):22-30;
1989.
10. Larsson, K. Conditioning and sexual behavior in the male albino rat.
In: Elmgren, I. J., ed. Acta psychologica
gothoburgensia.
Uppsala:
Almquist & Wiksells; 1956.
11. Larsson, K. Nonspecific stimulation and sexual behavior in the male
rat. Behaviour 20:110-114;
1963.
12. Longuski, P. A.; Cudillo, C. A.; Stern, J. J. Effects of estradiol on
feeding and locomotion in REMd rats. Physiol. Behav. 16:97-99;
1976.
13. Mark, J. Heiner, L. Mendel, P.; Godin, Y. Norepinephrine turnover
in brain and stress reactions in rats during paradoxical sleep deprivation. Life Sci. X:1085-1093; 1969.

14. Morden, B.; Mullins, R.; Levine, S. et al. Effects of REMS deprivation on the mating behavior of male rats. Psychophysiology
5:241242; 1968.
15. Rechtshaffen, A.; Bergmann, B. M.; Everson. C. A.; Kushida, C. A.;
Gilliland, M. A. Sleep deprivation in the rat: 1. Conceptual issues.
Sleep 12:1-4; 1989.
16. Rechtschaffen,
A.; Bergmann, B. M.; Everson, C. A.; Kushida, C.
A.; Gilliland, M. A. Sleep deprivation in the rat: X. Integration and
discussion of the findings. Sleep 12:68-87; 1989.
17. Retana-Marquez,
S.; Dominguez
Salazar,
E.; VelazquezMoctezuma, J. Effect of acute and chronic stress on masculine sexual
behavior in the rat. Psychoneuroendocrinology
(in press).
18. Sachs B, Meisel, R. The physiology of male sexual behavior. In:
Knobil, E.; Neil, J., eds. The physiology of reproduction. New York:
Raven Press. Ltd.: 1988:1393-1485.
19. Se&man, M. E. P. Helplessness: On depression, development and
death. San Francisco: Freeman; 1975.
20. Selye, H. The stress of life. New York: McGraw-Hill; 1976.
21. Stern, W. C.; Miller, F. P.; Cox, R. H.; Maickel, R. P. Brain
norepinephrine and serotonin levels following REM sleep deprivation
in the rat. Psychopharmacologia
22:50-55; 1971.
22. Velazquez-Moctezuma,
J.; Monroy, E.; Beyer, C.; Canchola, E.
Effects of REM deprivation on lordosis response induced by gonadal
steroids in ovariectomized rats. Physiol. Behav. 32:91-94; 1984.
23. Velazquez-Moctezuma,
J.; Monroy, E.: Cruz, M. L. Facilitation of
the effect of testosterone on male sexual behavior in rats deprived of
REM sleep. Behav. Neural Biol. 51:46-53; 1989.
24. Vimont-Vicary,
P. Jouvet, D.; Delorme, F. Effets EEG and comportementaux
des privations de sommeil paradoxale chez le chat.
EEG. Clin. Neurophysiol. 20:439-449; 1966.
25. Vogel, G. W. A review of REM sleep deprivation. Arch. Gcn.
Psychiatry 32:749-761;
1975.
26. Willner, P.; Muscat, R.; Papp, M. Chronic mild stress-induced anhedonia:a realistic animal model of depression. Neurosci. Biohehav.
Rev. 16:525-534;
1992.