See

discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/11587909

Naloxone blocks place preference conditioning

after paced mating in female rats

Article in Behavioral Neuroscience · January 2002

DOI: 10.1037/0735-7044.115.6.1363 · Source: PubMed

CITATIONS READS

81 19

2 authors, including:

Raúl G Paredes
Universidad Nacional Autónoma de México
104 PUBLICATIONS 2,772 CITATIONS

SEE PROFILE

All content following this page was uploaded by Raúl G Paredes on 04 November 2015.

The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document
and are linked to publications on ResearchGate, letting you access and read them immediately.
 Copyright 2001 by the American Psychological Association. Inc.
Behavioral Neuroscience 0735-7044/01/$5.00 DOJ: l0. l037//0735-7044. l l5.6. l363
2001. Vol. l 15. No. 6. 1363-1367

Naloxone Blocks Place Preference Conditioning After Paced Mating

in Female Rats

Raúl G. Paredes and Isadora Martínez

Universidad Nacional Autónoma de México

Sexual behavior in male rats induces a positive affect as evaluated by conditioned place preference
(CPP). In addition, when females control or "pace" the rate of sexual interaction, a clear CPP is also
observed. The reward state induced by mating in male rats is blocked by the injection of the opioid
antagonist naloxone. In the present experiment, a dose of 4 mg/kg of naloxone completely blocked the
CPP induced in females by paced mating. It appears that a common opioid system is involved in the
positive affect induced by sexual behavior in both male and female rats.

In rats, the possibility to control or "pace" the sexual interaction
is a crucial factor for sex to induce a positive affect. Males that
mated in a traditional testing chamber in which they controlled the
rate of sexual contacts developed a clear conditioned place pref-
erence (CPP; Agmo & Berenfeld, 1990; Martínez & Paredes, in
press). In addition, when females mated in a pacing chamber,
controlling the rate of sexual stimulation, they also developed a
clear CPP (Paredes & Alonso, 1997; Paredes & Vázquez, 1999).
Different hormone regimens can induce paced mating behavior
(Brandling-Bennett, Blasberg, & Clark, 1999), and, although there
are slightly different methodologies (for review, see Paredes &
Vázquez, 1999), the pacing chamber is usually divided by a wood
partition with a small hole. Because ofthe size ofthe hole, only the
female can enter or exit the half of the cage in which the male is
confined. In this methodology, the rats copulate in a mating cage
and are put in an adjacent conditioning chamber immediately after
sexual interaction (Paredes & Alonso, 1997; Paredes & Vázquez,
1999). Therefore, the affective state evaluated by CPP is not
limited to the execution of sexual behavior but is associated with
the physiological state induced by mating, allowing the association
of that state with environmental cues.
Severa) lines of evidence suggest that opioid peptides are in-
volved in sexual behavior (Agmo & Paredes, 1988). Researchers
using the CPP procedure have shown that sexual behavior
(Hughes, Herbert, & Everitt, 1990; Mehara & Baum, 1990; Miller
& Baum, 1987) and ejaculation (Agmo & Berenfeld, 1990) can
induce a reward state in male rats. This positive affect can be
blocked by the administration of the opiate antagonist naloxone
(Agmo & Gómez, 1993; Mehara & Baum, 1990; Miller & Baum,
Raúl G. Paredes and Isadora Martínez, Centro de Neurobiología, Uni-
versidad Nacional Autónoma de México (UNAM), Querétaro, Mexico.
This research was supported by Consejo Nacional de Ciencia y Tecno-
logía Grant 28039N and Dirección General de Asuntos del Personal
Académico Grant IN228 l 99. We thank Francisco Camacho and Lo urdes
Lara for technical assistance.
Correspondence concerning this article should be addressed to Raúl G.
Paredes, Centro de Neurobiología, UNAM, Apartado Postal 1-1141,
Querétaro, Querétaro 76001, Mexico. Electronic mail may be sent to
rparedes@servidor.unam.mx.
1987), suggesting that the release of opioids during sexual behav-
ior and ejaculation contribute to the rewarding consequences of
mating (reviewed in Agmo, 1999). In the present study, we eval-
uated whether the positive affect induced in female rats by paced
mating can be blocked by naloxone. This would let us determine
whether, as in males, opioids are involved in the rewarding con-
sequence of paced mating.
Method
Subjects
Sexually naive females (250-300 g) of the Wistar strain from a local
colony were maintained in a room with a reversed 12-hr light-dark cycle
(lights off at 0900). Commercial rat pellets (LabDiet, Nutrition Interna-
tional, Brentwood, MO) and water were always available. Subjects were
housed 2 per cage, depending on the group (see Design). Females were
bilaterally ovariectomized with a mixture of ketamine (95 mg/kg) and
xylazine (12 mg/kg). After surgery, each female received subcutaneous
injections of 25 µg estradiol benzoate, 48-52 hr before the mating test,
plus 1 mg progesterone 4 hr before testing. The steroids were dissolved in
corn oil and injected in a volume of 0.2 mi per rat. Males that ejaculated
in a 30-min test with receptive stimulus females were used as stimulus
males.
Apparatus
The place preference cage consisted of a three-compartment box made
ofwood. One ofthe lateral compartments (23 cm long X 37 cm high X 32
cm wide) had wood shavings on the floor and was painted white. The other
compartrnent was black, and the walls were moistened with a 2% solution
of glacial acetic acid before the subject was placed into it. In this way, the
lateral compartments offered distinct odor, color, and texture stimuli. The
middle compartment (22 cm long X 24 cm high X 32 cm wide) was
painted gray and communicated with the lateral compartments through a
sliding door (10 cm X 10 cm). The rats were observed through the front
wall of the middle compartment, which was made of fine wire mesh. The
place preference cages were located in a room illuminated with dim white
light. The mating cages (40 cm long X 60 cm high X 40 cm wide) were
located in the same room, adjacent to the preference cages. For paced
mating, the cages were divided by a removable wood partition with a small
hole, through which the female could enter or exit the half of the cage in
which the male was confined.

1363
1364 BRIEF COMMUNICATIONS
Drugs
Naloxone hydrochloride (Du Pont, Mexico) was dissolved in distilled
water and injected intraperitoneally, in a volume of l ml/kg. A dose of 4
mg/kg was injected 1 min before females were allowed to pace their sexual
interaction.
Design
Two groups were used. Group 1 females were injected with satine before
they were allowed to pace their coita! interactions, and Group 2 females
were injected with naloxone before they were allowed to pace their coita!
interactions. Both groups received identical hormonal treatment. They ali
were injected with estradiol benzoate 48 hr before mating and with pro-
gesterone 4 hr before mating (Conditioning Sessions 2, 4, and 6). Both
groups were drug free in the pretest and test sessions.
Procedure
In the pretest session, females were placed in the middle compartment,
and the time they spent in each of the lateral chambers during a 10-min
session was recorded. In this way, the initial preference was determined.
Subjects were assigned to one of two groups, corresponding to the time
spent in the preferred (nonreinforced) compartment. Six conditioning ses-
sions and a test followed the pretest. On the first, third, and fifth condi-
tioning session, ali subjects were injected with saline and transferred
directly from their home cage to the preferred compartment (nonrein-
forced) for 30 min without copulation. On alternate conditioning days, on
the second, fourth, and sixth session, Group 1 females were injected with
saline, and those of Group 2 were injected with naloxone. One minute after
injection, females were allowed to pace their sexual interaction. They were
placed in the left side of the mating cage, which was divided by the
removable wood partition. The mate was always confined to the right side.
Because of the small size of the hole, only the female could enter or exit
the half of the cage in which the male was confined. Immediately after
receiving an ejaculation, the females were gently withdrawn from the
mating test cage and placed individually in their corresponding nonpre-
ferred (reinforced) compartment for 30 min. An interval of 24-72 hr
between sessions was used, except for the interval between the last con-
ditioning session and the test, which was always 24 hr. After six condi-
tioning sessions, three reinforced and three nonreinforced, the preference
for each compartment was tested again in exactly the same way as before
conditioning (test session).
The following sexual behavior parameters were recorded: number of
mounts, intromissions, and lordosis displays. The lordosis intensity was
rated as follows: O = no lordosis displayed, 1 = medium lordosis after a
mount or an intromission, and 2 = foil lordosis displayed. From these
ratings, the mean lordosis intensity was calculated as the sum of points per
test/number of lordotic responses, and the lordosis quotient (LQ) was
calculated as (total number of lordosis responses / (total number of
mounts + intromissions) X 100). In addition, we calculated the mount
return latency (MRL; latency for the female to reenter the male's chamber
after a mount) and the intromission return latency (IRL; latency for the
female to return to the male's chamber after an intromission).
Statistical Analysis
Four criteria were used to evaluate place preference conditioning. The
first, time in the nonreinforced compartment, was expected to decrease
between pretest and test. The second, time in the reinforced compartment,
was expected to increase between pretest and test. The third was the
difference between time spent in the nonreinforced cage (initially preferred
si de) and time spent in the reinforced cage (initially nonpreferred si de
reinforced by paced mating). This measure had to be significantly reduced
after conditioning, and it eliminated the influence of the time spent in the
middle, neutral compartment. The fourth measure, preference score (time
in reinforced compartment / [time in reinforced compartment + time in
nonreinforced compartment]), was also used to minimize the possibility of
a spurious effect and was expected to increase between pretest and test.
Each of the preference measures were evaluated by a 2 (test) X 2 (group)
analysis of variance (ANOV A) with repeated measures on the test factor
(pretest-test). The sexual behavior parameters were evaluated by a 3
(conditioning session) X 2 (groups) ANOV A with repeated measures on
the session factor. The female's MRL and IRL were not analyzed by a
two-way repeated measures ANOV A because there were many missing
values. These were the result of males not mounting or females not exiting
after a mount oran intromission, thus making this analysis impossible. The
MRL was analyzed by a Kruskal-Wallis one-way ANOV A because of the
reduced number of observations. The IRL was analyzed by a two-way
ANOVA.
Results
Place Preference
The ANOV A for the time spent in the nonreinforced compart-
ment revealed an effect of session, F(l, 23) = 10.48, p = .004;
group, F(l, 23) = 1.08, p = .31; and a Group X Session interac-
tion, F(l, 23) = 9.43, p = .005. Tests for simple main effects
showed a significant reduction in the time spent in the nonrein-
forced compartment in females that paced their sexual contacts
after saline injection, F(l, 23) = 19.14, p < .001. No change in
preference was observed in pacing females treated with naloxone,
F(l, 23) = O.O!, p = .90.
A significant Group X Session interaction, F( 1, 23) = 34.26, p
< .001, and nonsignificant effects of session, F(l, 23) = 1.96, p =
.17, and group, F(l, 23) = 0.09, p = .76, were observed in the time
spent in the reinforced compartment. Tests for simple main effects
showed that the group treated with saline significantly increased
the time spent in the reinforced compartment, F(l, 23) = 25.29, p
< .001. An opposite effect was observed in the group treated with
naloxone, that is, the time in the reinforced compartment decreased
between pretest and test, F{l, 23) = 10.33, p = .004.
The analysis of the difference between compartments revealed a
significant effect of session, F(l, 23) = 10.38, p = .004, and
Group X Session interaction, F( l, 23) = 33.53, p < .001, but no
significant effect of group, F(l, 23) = 0.74, p = .39. Tests for
simple main effects were then performed. The difference between
time spent in the nonreinforced compartment and time spent in the
reinforced compartment was significantly reduced in the group of
females injected with saline after pacing their sexual contacts, F(l,
23) = 39.05, p < .001. No reduction in the difference between
time spent in the nonreinforced compartment and time spent in the
reinforced compartment was observed in the group of females
treated with naloxone before they paced their sexual interaction,
F(l, 60) = 3.44, p = .07.
The ANOV A of the preference score revealed a significant
effect of session, F(l, 23) = 14.33, p = .001, and Group X
Session interaction, F(l, 23) = 37.47, p < .001, but no significant
effect of group, F(l, 23) = 0.79, p = .38. Test for simple main
effects showed a significant increase in the preference score in the
group of females injected with saline before they paced their
sexual interaction, F(l, 23) = 47.18, p < .001. No change in the
preference score was observed in the females injected with nalox-
one before they paced their sexual interaction, F(l, 23) = 2.84, p
= . l O. Figure 1 illustrates the clear preference induced by paced
 BRIEF COMMUNICATIONS 1365

D PRETEST ~ TEST
E- 250
A B ***
;z: E-
u.:¡ ;z: 160
¿: u.:¡
E- ¿:
o::
<C 200
o..
¿:
o
E-
o::
<C
o.. 120
%//
·0/,////
//
u *** ;E
o / ///• **
ou.:¡ 150 u /;;,;::/.·
.~/·
u ou.:¡ // / //, / /, /
o:: /,
/

<~
ou. u 80 /,
///
o::
ou. <::/:/,/, /

/;
;z: 100 ////.·. /
¡;j //// ;z: //. // ///•
o:: //: / /
/:.// ¡;j ' //; ; '/ // / ///·'
z
o;z:
·>/~
·.//// o:: 40
'/

/><,>;··,,
/
.' /
/

·,~)
50 2!::
//~ u.:¡ /'/:>/ /
2!::
u.:¡
¿:
".' /
•/·////
¿:
¡::
/ //.
//;/~///
// / / /
/
/<' ;;·
// //
¡:: o
/,./ /
o

e 0.7 D
"'E- 150 ***

~
0.6

0.5
<C
o.. 100
¿:
o ~
u o 0.4
/ / /

u
"'u
~
u.:¡ /
/
50 0.3
E-
u.:¡
al
~
u.:¡
•//

u.:¡
u o
*** , / ""~ 0.2 /

~
u.:¡
0.1

s
""
-50 00

SALINE NALOXONE SALINE NALOXONE

Figure J. Time in the nonreinforced compartrnent (A), time in the reinforced compartment (B), difference
between time spent in the nonreinforced and reinforced compartrnent (C), and preference score (D), at pretest
and test Subjects were mated in a charnber divided by a wood partition with a small hole, through which only
the female could enter or exit the half in which the male was confined. Females were treated with saline or
naloxone (4 mglkg) 1 rnin before being allowed to pace their sexual interaction. Data are expressed as means
(::+:: SEMs). Time is expressed in seconds. Asterisks indicate a significan! difference from pretest (**p < .01,
***p < .001).

mating when the females were injected with saline. The injection
of naloxone completely blocked the change in preference induced
by paced mating.
Sexual Behavior
When the mean lordosis intensity (MLI) was analyzed, no
significant differences were found: group, F(l, 23) = 1.92, p =
.18; session, F(2, 46) = 0.91, p = .41; Group X Session, F(2,
46) = 0.70, p = .50. Moreover, no significant differences were
observed in the LQ. In fact, because ali females were receptive in
ali the sessions, they had an LQ of 1OO. Because there was no
variance, the ANOV A was not performed. These results indicate
that ali females were equally receptive throughout conditioning
(see Table 1).
The analysis of the number of mounts that the females
received revealed no significant differences: group, F(l,
23) = 0.31, p = .58; session, F(2, 46) = 0.80, p = .45;
Group X Session, F(2, 46) = 1.21, p = .30. Similarly, no
differences were observed in the number of intromissions re-
ceived: group, F(l, 23) = 0.29, p = .59; session, F(2,
46) = 2.05, p = .14; Group X Session, F(2, 46) = 0.46, p =
.63. No significant differences were observed in the MRL (p =
.03) or the IRL: group, F(l, 34) = 0.96, p = .34; session, F(2,
1366 BRIEF COMMUNICATIONS

Table 1
Sexual Behaviors in Fema/e Rats lnjected With Satine or Naloxone (4 mglkg) 20 Min Befare
They Paced the Sexual lnteraction in Three Mating Sessions

Satine and pacing Naloxone and pacing

Behavior 2 3 2 3

MLI 1.3 ::t: 0.9 1.3 ::t: 0.8 1.3 ::t: 0.9 1.5 ::t: 0.9 1.3 ::t: 0.8 1.4 ::t: 0.9
LQ IOO IOO IOO IOO IOO IOO
M 3.5 ::t: 0.9 3.0 ::t: 1.3 2.4 ::t: 0.8 1.8 ::t: 0.8 1.8 ::t: 0.6 9.7 ::t: 7.5
I 9.3 ::t: 0.8 10.1 ::t: 0.8 12.6 ::t: 1.7 8.4 ::t: 0.9 10.3 ::t: 2.0 14.2 ::t: 6.9
MRL 12.5 ::t: 9.2 15.4 ::t: I0.6 23.7 ::t: 16.5 3.7 ::t: 2.6 33.6 ::t: 19.9 6.6 ::t: 4.5
IRL 33.9 ::t: 13.8 92.9 ::t: 32.8 90.0 ::t: 26.4 101.6 ::t: 21.9 103.7 ::t: 33.1 86.0 ::t: 21.6

Note. Ali data (except lordosis quotient [LQ]) are expressed as means (::t: SEMs). Mount retum latency (MRL)
and intromission retum latency (IRL) are expressed in seconds. MLI = mean lordosis intensity; M = number
of mounts received; I = number of intromissions received.

34) = 1.70, p = .19; Group X Session, F(2, 34)
. 17. These data are summarized in Table 1.
Discussion
The group of females injected with saline throughout condition-
ing developed CPP, further supporting the suggestion that paced
coita! contacts induce a positive affect of sufficient intensity and
duration to induce conditioning (Paredes & Alonso, 1997; Paredes
& Vázquez, 1999). lt is well established that the release of opioids
during sexual behavior and ejaculation contribute to the rewarding
consequences of mating in male rats (Agmo & Berenfeld, 1990;
Agmo & Paredes, 1988; Hughes et al., 1990; Mehara & Baum,
1990; Miller & Baum, 1987), and that this positive, mating-
induced affect can be blocked by the administration of the opiate
antagonist naloxone (Agmo & Gómez, 1993; Mehara & Baum,
1990; Miller & Baum, 1987). In the present study, the injection
of 4 mg/kg naloxone completely blocked CPP induced in females
by paced mating. This dose has been previously used to block
incentive motivational responses to opioid-related stimuli evalu-
ated by CPP (Agmo & Berenfeld, 1990; Paredes, Muzzi, Aguirre,
& Romero, 2000). It has been suggested that opioid release during
sexual behavior contributes to the elimination of aversive conse-
quences of mating, possibly as a result of the analgesic effect of
copulation. For example, analgesic effects have been described as
occurring both before ejaculation (Szechtman, Hershkowitz, &
Simantov, 1981) and after ejaculation (Forsberg, Wiesenfeld-
Hallin, Eneroth, & Sodersten, 1987) in male rats, as well as after
vaginal stimulation in females (Komisaruk & Wallman, 1977).
The administration ofthe opioid antagonist completely blocked the
rewarding properties of paced mating. Although we did not di-
rectly test the possibility that the injection of naloxone per se
induced place aversion, previous studies using exactly the same
paradigm have shown that this is not the case (Agmo & Berenfeld,
1990).
Mehara and Baum (1990) showed that, although naloxone failed
to affect the acquisition of CPP for a female in estros, the opioid
antagonist strongly blocked the expression of an established CPP
in males. That is, administering naloxone to male rats before their
mating with a receptive female failed to block CPP. In contrast,
when males were injected with saline during conditioning and
= 1.85, p = injected with naloxone before the final tests, CPP was blocked.
The authors interpreted their results as suggesting that CPP occurs
in the absence of primary rewarding stimuli but depends on the
conditioned incentive stimuli previously associated with mating.
These observations appear to be at variance with those reported by
Agmo and Berenfeld (1990), in which naloxone was able to block
the acquisition of CPP in male rats after ejaculation. The results of
the present experiment are in agreement with those reported by
Agmo and Berenfeld (1990); that is, the female received naloxone
injection before mating with amale rat, suggesting that the acqui-
sition of CPP was blocked. However, one important difference is
that, whereas in the Mehara and Baum (1990) study mating oc-
curred in the conditioning cage, in Agmo and Berenfeld' s (1990)
study and in our own work subjects were mated in an adjacent
mating cage and were placed in the conditioning cage immediately
afterward. In this latter case, the positive affect induced by copu-
lation is not associated with the execution of sexual behavior itself,
but rather with the physiological state induced by mating. lt could
be argued, then, that in three studies naloxone did not block the
primary rewarding stimuli (i.e., copulation itself), but rather the
conditioned incentive stimuli or the physiological state induced by
mating.
An increase in doparnine (DA) release in the striatum and
nucleus accumbens has been described in females that paced their
sexual contacts in comparison with nonpacing females (Mermel-
stein & Becker, 1995), indicating that DA neurotransmission
might be involved in paced mating (Mermelstein & Becker, 1995;
Pfaus, Damsma, Wenkstem, & Fibiger, 1995). Moreover, the CPP
induced by mating in the female Syrian hamster, which does not
pace sexual interactions, was blocked by the DA 2 receptor antag-
onist raclopride (Meisel, Joppa, & Rowe, 1996). However, another
DA antagonist, pimozide, was not able to block ejaculation-
induced reward in male rats (Agmo & Berenfeld 1990). Further
studies are needed to determine whether DA antagonists can block
the positive affect induced by paced mating in female rats. The
results of the present experiment clearly indicate that naloxone
effectively blocks CPP induced by paced mating, which is prob-
ably produced by the release of endogenous opioids. These results,
together with those described in male rats, suggest that a common
opioid system mediates sexual reward in male and female rats.
 BRIEF COMMUNICATIONS
References
Agmo, A. (1999). Sexual motivation: An inquiry into events determining
the occurrence of sexual behavior. Behavioural Brain Research, 105,
129-150.
Agmo, A., & Berenfeld, R. ( 1990). Reinforcing properties of ejaculation in
the male rat: Role of opioids and dopamine. Behavioral Neuroscience,
104, 177-182.
Agmo, A., & Gómez, M. (1993). Sexual reinforcement is blocked by
infusion of naloxone into the medial preoptical area. Behavioral Neu-
roscience, 107, 812-818.
Agmo, A., & Paredes, R. (1988). Opioid and sexual behavior in the male
rat. Pharmacology Biochemistry and Behavior, 30, 1021-1034.
Brandling-Bennett, E.M., Blasberg, M. E., & Clark, A. S. (1999). Paced
mating behavior in female rats in response to different hormone priming
regimens. Hormones and Behavior, 35, 144-154.
Forsberg, G., Wiesenfeld-Hallin, Z., Eneroth, P., & Sodersten, P. (1987).
Sexual behavior induces naloxone-reversible hypoalgesia in male rats.
Neuroscience Letters, 81, 151-154.
Hughes, A.M., Herbert, J., & Everitt, B. J. (1990). Comparative effects of
preoptic area infusions of opioid peptides, lesions and castration on
sexual behavior of male rats: Studies of instrumental behavior, condi-
tioned place preference and partner preference. Psychopharmacology
(Berlin), 102, 243-256.
Komisaruk, B. R., & Wallman, J. (1977). Antinociceptive effects of vag-
inal stimulation in rats: Neurophysiological and behavioral studies.
Brain Research, 137, 85-107.
Martínez, I., & Paredes, R. G. (in press). Only self-paced mating is
rewarding in rats of both sexes. Hormones and Behavior.
Mehara, B. J., & Baum, M. J. (1990). Naloxone disrupts the expression but
not the acquisition by male rats of a conditioned place preference
1367
response for an oestrus female. Psychopharmacology (Berlin), 101,
118-125.
Meisel, R. L., Joppa, M. A., & Rowe, R. K. (1996). Dopamine receptor
antagonists attenuate conditioned place preference following sexual be-
havior in female Syrian hamsters. European Journal of Pharmacology,
309, 21-24.
Mermelstein, P. G., & Becker, J. B. (1995). Increased extracellular dopa-
mine in the nucleus accumbens and striatum of the female rat during
paced copulatory behavior. Behavioral Neuroscience, 109, 354-365.
Miller, R. L., & Baum, M. J. (1987). Naloxone inhibits mating and
conditioned place preference for an estrous female in male rats soon
after castration. Pharmacology Biochemistry and Behavior, 26, 781-
789.
Paredes, R. G., & Alonso, A. ( 1997). Sexual behavior regulated (paced) by
the female induces conditioned place preference. Behavioral Neuro-
science, 111, 123-128.
Paredes, R. G., Muzzi, G., Aguirre, E., & Romero, V. (2000). Can a
generalized kindling seizure induce a reward state? Epilepsy Research,
38, 249-257.
Paredes, R. G., & Vázquez, B. (1999). What do female rats Iike about sex?
Paced mating. Behavioural Brain Research, 105, 117-127.
Pfaus, J. G., Damsma, G .. Wenkstern, D., & Fibiger, H. C. (1995). Sexual
activity increases dopamine transmission in the nucleus accumbens and
striatum of female rats. Brain Research, 693, 21-30.
Szechtman, H., Hershkowitz, M., & Simantov, R. (1981). Sexual behavior
decreases pain sensitivity and stimulates endogenous opioids in male
rats. European Journal of Pharmacology, 70, 279-285.
Received October 6, 2000
Revision received February 22, 2001
Accepted May 10, 2001 •
 Instructions to Authors
Behavioral Neuroscience
Behavioral Neuroscience is a bimonthly, peer-reviewed
journal that publishes original research articles in the broad
field of the biological bases of behavior. A detailed descrip-
tion of the editorial coverage policy appears on the inside of
the front cover of each issue.
Manuscript Preparation
Authors should prepare manuscripts according to the
Publication Manual of the American Psychological Associ-
ation (4th ed.). Typing instructions (all copy must be
double-spaced with wide margins on 8.5- X 11-inch paper)
and detailed instructions on preparing abstracts, tables, fig-
ures, references, and abbreviations and metrics appear in the
Publication Manual. Also, all manuscripts are copyedited
for bias-free language. The journal also publishes a ''Brief
Communications'' section. Articles for this section should
be so labeled and must not exceed 3,250 words of text, with
no more than two figures and/or tables. The text count <loes
not include references or figure captions. ''Technical Com-
ments'' concerning articles previously published in the jour-
nal will also be considered for publication. (Articles for this
section must not exceed 800 words of text.)
Abstracts
All manuscripts must include an abstract that contains a
maximum of 960 characters and spaces (which is approxi-
mately 120 words) typed on a separate sheet of paper.
Tables
Each table should be submitted on a separate page and
must be numbered and labeled with an appropriate title. All
tables must be self-explanatory.
Figures
To ensure high-quality reproduction of photomicro-
graphs, each issue of Behavioral Neuroscience, as well as
any reprints of articles that authors purchase, will be printed
on coated, photographic-quality paper. To further ensure the
quality of all figures in the journal, authors should note that
figures must be photographed and submitted as high-quality
glossy prints or laser prints, which must not exceed 8.5 X
11 inches (21.5 X 28 cm) in size; photocopies, transparen-
cies, slides, and negatives are not acceptable. Photomicro-
graphs must not be mounted on cardboard or poster board,
and all parts of multiple-part figures must appear on the
same glossy print. All figures must be numbered and labeled
with the first author's name. Original color figures can be
printed in color provided the author agrees to pay half of the
associated production costs.
References
References should be cited in text as follows: "The
results replicated those of a previous study (Knoth & Mair,
1991 ), " or "The procedure was a modification of Krettek
and Price (1989) and Smith et al. (1977) .... " Multiple
references should be cited in alphabetical order: "Earlier
investigations (Abbott, 1988; Hunt & Aggleton, 1983; Win-
ocur, 1985).... '' Each listed reference should be cited in
1368
text, and each text citation should be listed alphabetically in
the reference section. The following examples illustrate the
style to be used for a journal article, a book, and a chapter
in a book, respectively:
Crider, A. B., Blockel, L. L., & Solomon, P. R. (1988).
Selective attention in the Fisher 344 rat: Dopamine receptor
supersensitivity or up-regulation? Behavioral Neuroscience,
102, 315-319.
Paxinos, G., & Watson, C. (1986). The rat brain in
stereotaxic coordinates (2nd ed.). San Diego, CA: Aca-
demic Press.
Zola-Morgan, S. (1984). Toward an animal model of
human amnesia: Sorne critica} issues. In L. R. Squire & N.
Butters (Eds.), Neuropsychology of memory (pp. 83-103).
New York: Guilford Press.
Abbreviations and Metrics
Nonstandard abbreviations should be introduced by plac-
ing the abbreviation in parentheses after the first occurrence
of the term being abbreviated in both the abstract and the
text. The metric system should be followed for all volumes,
lengths, weights, and so on. Temperatures should be ex-
pressed in degrees celsius (centigrade). Units should con-
form to the International System of Units (SI; see the
Publication Manual).
Manuscript Submission
Manuscripts should be submitted in quadruplicate, and all
copies should be clear, readable, and on paper of good
quality. A dot matrix or unusual typeface is acceptable only
if it is clear and legible. In addition to addresses and phone
numbers, authors should supply electronic mail addresses
and fax numbers, if available, for potential use by the
editorial office and later by the production office. Upon
acceptance of their manuscript, authors will also be re-
quested to submit an electronic version of their manuscript
on diskette. Authors should keep a copy of their manuscript
to guard against loss. Effective in January 2001, the Incom-
ing Editor is receiving all new submissions to the journal.
Submissions that are accepted will be published beginning
in the 2002 volume. Mail manuscripts to John F. Disterhoft,
Incoming Editor, Behavioral Neuroscience, Department of
Cell and Molecular Biology, Northwestern University Med-
ica! School, Ward Bldg., Rm. 5-003, 303 East Chicago
Avenue, Chicago, IL 60611.
Submission Letter
Authors are required to include the following in their
submission letter:
1. A statement of compliance with APA ethical stan-
dards in the treatment of their sample, human or animal, or
a description of the details of the treatment.
2. A statement that the manuscript or data have not been
 INSTRUCTIONS TO AUTHORS
published previously and that they are not under consider-
ation for publication elsewhere.
3. A statement to reflect that all listed authors have
contributed significantly to the manuscript and consent to
their names on the manuscript.
4. A statement to disclose any possible conflict of inter-
est in the conduct and reporting of research (e.g., financia!
interests in a test or procedure, funding by pharmaceutical
companies for drug research).
Authors are encouraged to suggest five reviewers who are
especially qualified to review their work and would not
have a conflict of interest serving as a referee.
Publication Policy
Authors are required to obtain and provide to APA all
necessary permissions to reproduce any copyrighted work,
including, for example, test instruments and other test ma-
terials or portions thereof.
APA policy prohibits an author from subrnitting the same
manuscript for concurrent consideration by two or more
publications. In addition, it is a violation of AP A Ethical
Principies to publish "as original data, data that have been
previously published" (Standard 6.24). Because this joumal
is a primary joumal that publishes original material only,
APA policy prohibits as well publication of any manuscript
that has already been published in whole or substantial part
elsewhere. Authors have an obligation to consult joumal
editors conceming prior publication of any data upon which
their article depends. In addition, APA Ethical Principies
specify that "after research results are published, psychol-
ogists do not withhold the data on which their conclusions
are based from other competent professionals who seek to
verify the substantive claims through reanalysis and who
intend to use such data only for that purpose, provided that
the confidentiality of the participants can be protected and
unless legal rights conceming proprietary data preclude
their release" (Standard 6.25). APA expects authors sub-
rnitting to this joumal to adhere to these standards. Specif-
ically, authors of manuscripts subrnitted to APA joumals
are expected to have available their data throughout the
editorial review process and for at least 5 years after the date
of publication.
Review Policy
Masked reviews are optional, and authors who wish
masked reviews must specifically request them when sub-
rnitting their manuscripts. Each copy of a manuscript to be
mask reviewed should include a separate title page with
authors' names and affiliations, and these should not appear
anywhere else on the manuscript. Footnotes that identify the
authors should be typed on a separate page. Authors should
make every effort to see that the manuscript itself contains
no clues to their identities.
View publication stats
1369
Revisions
Revised manuscripts and all correspondence should be
sent to the main editorial office at Johns Hopkins Univer-
sity. Manuscripts need not be accompanied by a copy of the
original version. Revisions not retumed within two months
of the last action date will be treated as a new subrnission.
Manuscript Acceptance
Upon acceptance of their manuscript for publication, au-
thors are expected to provide permissions, signed and dated
copyright release and ethical compliance forms, and an
electronic version of their revised manuscript on diskette.
Permissions
If any figure or table (or part or adaptation thereof) or
more than a few lines of text from previously published
material are included in a manuscript, the author must
obtain written permission for re-publication from the copy-
right holder and forward a copy to the AP A Joumals Office,
750 First Street, NE, Washington, DC 20002-4242.
Copyright
Under copyright law, the transfer of copyright from au-
thor to publisher must be completed befare any article can
be published in Behavioral Neuroscience. The transfer of
copyright enables the publisher to assure maximum dissem-
ination of the author' s work. Copyright forms are sent to all
authors prior to acceptance and must be signed and retumed
to the Editor's office immediately. U.S. govemment em-
ployees must sign the section of the form stating exemption
from copyright laws. Alterations to or substitutions for the
form are not acceptable.
Ethical Compliance
Authors will be required to state in writing that they have
complied with APA ethical standards in the treatment of
their sample, human or animal, orto describe the details of
treatment. A copy of the APA Ethical Principies may be
obtained by writing the AP A Ethics Office, 750 First Street,
NE, Washington, DC 20002-4242.
Proofs and Reprints
All proofs must be corrected and retumed to the APA
Joumals Office within 48 hours of receipt. Foreign authors
are encouraged to use a rapid and reliable mailing service.
Only correction of typographical errors will be permitted;
the author will be charged for additional alterations to the
text at the proof stage.
With the proofs will be a form providing the author with
the opportunity to order reprints. Direct inquiries to the
APA Joumals Office can be made at 202-336-5540; fax
202-336-5549.