Effects of taurine on rat behaviors in three anxiety models
Wei Xi Kong1, Si Wei Chen⁎, Yu Lei Li, Yi Jing Zhang, Rui Wang, Li Min2, Xiaojuan Mi
Department of Pharmacology, Shenyang Pharmaceutical University, Box 41, 103 Wenhua Road, 110016 Shenyang, P. R. China
Received 10 May 2005; received in revised form 26 January 2006; accepted 1 February 2006
Available online 15 March 2006
In our previous studies using an elevated plus-maze test in mice, taurine was shown to present an anxiolytic-like effect after single and repeated
administration [Chen SW, Kong WX, Zhang YJ, Li YL, Mi XJ, Mu XS. Possible anxiolytic effects of taurine in the mouse elevated plus-maze.
Life Sci 2004;75: 1503–11]. The aim of the present study was to investigate the anxiolytic and behavioral effects of taurine on rats in the open
field, hole-board, and social interaction test compared to the positive control diazepam. Taurine (14, 42, and 126 mg/kg, i.p.) was administered
30 min before the tests. In the social interaction and hole-board tests, taurine (42 mg/kg) significantly increased social interaction time and the
number and duration of head-dipping. In the open field test, taurine (126 mg/kg, i.p.) presented anxiolytic-like effects by increasing the number of
center entries, time spent in the central area and the anti-thigmotactic score while having no effect on the locomotor activity. Results from these
experiments suggest that taurine produces an anxiolytic-like effect in these animal models and may act as a modulator or anti-anxiety agent in the
central nervous system.
© 2006 Elsevier Inc. All rights reserved.
Keywords: Taurine; Anxiolytic; Diazepam; Social interaction; Hole-board; Open field; Rat
Taurine is one of the most abundant amino acids in the
brain (Jacobsen and Smith, 1968). The high levels of taurine in
this organ have stimulated much research to establish the
possible function for it. The role of taurine as a putative
neurotransmitter was reviewed by Kuriyama (1980). McBride
and Frederickson (1979) proposed taurine as a possible
inhibitory transmitter in the cerebellum. Taurine levels are
increased during stress, hypoxia, energy deprivation (Mila-
kofsky et al., 1984; Bockelmann et al., 1998; Colivicchi et al.,
1998) and regions that have extremely high taurine levels or
are very sensitive to taurine manipulation include hippocampus
(Galarreta et al., 1996), striatum (Lombardini, 1977), and
corticost riatal proje ction ( Sergeeva a nd Haas, 2001 ). Fo llow-
ing systemic administration taurine enters the brain via a
sodium and chloride dependent carrier from blood to the
endothelial cell (Benrabh et al., 1995; Molchanova et al.,
2004). Endogenous taurine is released from nervous tissue in
response to depolarizing agents such as N-methyl-D-aspartate
(NMDA) and high concentration of K+(Saransaari and Oja,
2003) and sequestered by an active high-affinity uptake system
(Oja and Kontro, 1984). Extracellular taurine modifies the
release of amino acid transmitters and modulates intracellular
Ca2+homeostasis (Foos and Wu, 2002). Although the mode of
action of taurine still remains to be elucidated, it has been
shown that taurine affects the metabolism of transmitters such
as γ-aminobutyric acid (GABA) (Medina and De Robertis,
1984; Kontro and Oja, 1990; Michel and Richard, 1991;
Liljequist, 1992) and 5-hydroxytryptamine (5-HT) (Sgaragli et
al., 1981; Becquet et al., 1993). Taurine inhibits the Ca2+-
dependent release of GABA and reduces 5-HT concentration
in the hypothalamus. In the rat striatum, strychnine-sensitive
glycine receptors are present on cholinergic interneurons
(Darstein et al., 2000) and taurine, together with glycine, is
their highly potent agonist (Sergeeva and Haas, 2001).
Behaviorally, i.p. injections of taurine (0.3–3.0 mg/kg) have
produced a dose-dependent depression of habituated psychomotor
Pharmacology, Biochemistry and Behavior 83 (2006) 271–276
⁎Corresponding author. Tel.: +86 24 23909448; fax: +86 24 24513031.
E-mail address: email@example.com (S.W. Chen).
1Present address: Department of Pharmacology, Dalian Medical University,
465 Zhongshan Road, 116027 Dalian, P. R. China.
2Present address: Department of phytochemistry, College of Pharmacy,
Second Military Medical University of PLA, 800 Xiangyin Road, 200433
Shanghai, P. R. China.
0091-3057/$ - see front matter © 2006 Elsevier Inc. All rights reserved.
activity in rats (Baskin et al., 1974). Sanberg and Ossenkopp (1977)
reported that increasing dose (25–200 mg/kg) of taurine signif-
icantly decreased ambulation levels, increased latency scores and
increased thigmotaxis in the rat open field test. In mice, i.p.
injections of large amounts (9–21.3 mmol/kg) of this amino acid
resulted in decreased locomotor activity and decreased instrumental
responding for food or water (Hruska et al., 1975). In our previous
studies using an elevated plus-maze test in mice, taurine could
significantly increase the percentage of time in the open arms after
acute oral administration (60 mg/kg) and repeated oral administra-
that taurine might play a role in the modulation of anxiety. In the
present study, we were interested to see whether it will exhibit
similar effect in the same kind of models in rats. We selected three
The social interaction test provided a good model of generalized
anxiety disorder (GAD) (Cheeta et al., 2000). The test conditions
were manipulated to generate different levels of anxiety and both
anxiolytic and anxiogenic drug effects could be detected. The hole-
boardtest andopen-fieldtest provide simple methods for measuring
used to assess emotionality, anxiety and/or responses to stress in
animals. Diazepam is still the most widely used and an effective
anxiolytic drug, and it has been found to be effective in many
anxiety models including society interaction, hole-board and open
field test (Kamei et al., 2001; Min et al., 2005). So, diazepam was
chosen as the positive control in our present study. The doses of
mice in our previous studies were converted to the equivalent rat
dose based on body surface area.
2. Material and methods
Male Wistar rats (Experimental Animal Center of Shenyang
Pharmaceutical University) weighing 180–200 g were kept
under a 12 h reversed light cycle (light off 07:00) at 22±2 °C
with free access to food and water for at least 7 days before
experimentation. During this period they were handled daily
and the position of the cages in the rack was changed so that all
rats received equal experience of the different levels of illu-
mination. In the social interaction test rats were individually
housed (cage size: 25×14×12 cm) and in the hole-board and
open field test rats were housed in group of five. Independent
animals were tested in each of these paradigms and in all tests of
anxiety, rats were injected in their holding rooms before testing
in an adjacent laboratory.
All animal treatments were strictly in accordance with the
National Institutes of Health Guide of the Care and Use of
Laboratory Animals. The experiments were carried out under
the approval of the Committee of Experimental Animal
Administration of the University.
2.2. Drugs and treatments
Taurine, 2-aminoethane-sulphonic acid, was purchased from
Shanghai Reagent Co. (Shanghai, China), diazepam from Hubei
Pharmaceutical Factory (Hubei, China), and Tween 80 from
Shenyang Dongxing Reagent Factory (Shenyang, China).
Diazepam (2 mg/kg for social interaction and open-field test
and 0.3, 0.6 mg/kg for hole-board test) and taurine (14, 42,
126 mg/kg) were suspended by ultrasound in 0.9% saline to
which Tween 80 (2 drops/10 ml) had been added. All drugs
were prepared freshly on test days and administered i.p. in a
volume of 2 ml/kg 30 min before testing. Control animals were
administered with the vehicle.
3. Behavioral tests
3.1. Social interaction test
The general design was essential as reported by File and
Hyde (1978). The test was conducted in a Perspex box with
opaque walls on four sides (60×60×35 cm), the floor of which
was divided into nine (20×20 cm) squares. The test conditions
were manipulated by altering the familiarity of rats to the test
arena. Two test conditions were performed: high light level in
unfamiliar conditions (HU) and in familiar conditions (HF). The
light intensity of the arena was 380 lux.
A total of 60 male rats were divided into five treatment
groups: vehicle control, diazepam (2.0 mg/kg), TA (14, 42 and
126 mg/kg). At the first day of the test, rats were tested in the
HU condition. Each rat was tested for social interaction with an
unknown test partner that did not differ by more than 15 g in
weight. Both members of a pair had the same prior
familiarization experience and the same drug treatment. Pairs
of rats were placed in opposite corners of the arena and then left
for 10 min. Their behaviors were recorded with a video camera
and observed on a monitor in an adjacent room. The total time
of non-aggressive, active social interactions including sniffing,
nipping, allogrooming, following, jumping on, crawling under
and over the partner and locomotor activity (the number of
squares crossed) behaviors were recorded for each pair by two
blind observers and the average scores used for subsequent
analysis. Passive body contact was not regarded as a social
interaction. After the first day of test, rats were returned to their
home-cages. In the following two consecutive days, these rats
were placed singly, undrugged, in the same test box for 10 min
to familiarize them with the environment. On the fourth day, the
same pairs of rats were once again tested in the HF condition
and the same test procedure was carried out.
3.2. Hole-board test
The hole-board apparatus consisted of Perspex box
(60×60×35 cm) with four equidistant holes 4 cm in diameter
in thefloor.The floor of the box was positioned 12 cm above the
ground and divided into nine (20×20 cm) squares. For the hole-
board experiments, each animal was placed in the center of the
hole-board and allowed to freely explore the apparatus for
5 min. Total number of squares crossed, number and duration of
rearing and head-dipping, and latency to the first head-dipping
were recorded by a video camera. Videotapes were later scored
by a trained observer blind to the treatment conditions.
272W.X. Kong et al. / Pharmacology, Biochemistry and Behavior 83 (2006) 271–276
3.3. Open-field test
A round (80-cm diameter) open field with an opaque black
floor surrounded by 30-cm high walls was used for the
behavioral test. The floor was marked off in two concentric
circles (the inner circle's diameter is 50 cm) and divided into
10×10 cm squares. Four 25-Wred bulbs 100 cm above the field
provided illumination and the test was performed in a quiet
room without previous habituation. The experimental sessions
were recorded by a video camera interfaced with a monitor and
a videocassette recorder in an adjacent room. Each animal was
placed in the periphery of the arena and three behavioral
measures were recorded in the 5-min test duration: locomotor
activity (the number of squares entered by the four paws of the
rat), the number and duration of center entries (defined as a
movement of an animal from the wall to the central area crossed
the inner circle line). The anti-thigmotactic effect was calculated
as a ratio of the number of entries into the central part to the
locomotor activity and multiplied by 1000 (Siemi1tkowski et
al., 2000). A higher value in score indicates a more pronounced
anxiolytic-like effect. This parameter was calculated for each rat
separately, and then the mean value for each experimental group
Results are reported as means±SEM. Data were analyzed by
means of analysis of variance (ANOVA). Whenever ANOVA
was significant, further multiple comparisons were made using
the Dunnett's t-test. All analyses were performed using the
software SPSS V11.5 for windows. The level of statistical
significance adopted was Pb0.05.
4.1. Social interaction test
The results for the social interaction test are shown in Fig. 1.
In the HU condition there was a significant drug-induced
increase in social interaction [F (4,25)=4.02, Pb0.05]. Further
analyses confirmed that both diazepam (2 mg/kg) and TA
(42 mg/kg) significantly increased social interaction time
compared with the control group (Pb0.01). In the HF condition
there was again a significant drug-induced increase in social
interaction [F (4,25)=5.31, Pb0.05], due to the dose of 42 mg/
kg of TA (Pb0.05), although diazepam (2.0 mg/kg) had no
effect on the total time spent in social interaction. Both taurine
social interaction time (s)
TA (mg/kg)TA (mg/kg)
Fig. 1. Effects of taurine and diazepam on total interactiontime and locomotion (numberof squares crossed) in a 10-min social interaction test. Results given as mean±
SEM (n=5∼6 pairs). Pair of rats was treated as a unit and tested 30 min after i.p. of vehicle, taurine (14, 42, 126 mg/kg) and diazepam (2 mg/kg). Only one score for
the pair was used.⁎Pb0.05,⁎⁎Pb0.01 compared with vehicle group. HU: high light, unfamiliar condition; HF: high light, familiar condition.
Effects of taurine and diazepam on exploratory behavior in rats tested on the hole-board test
Number of squares
Data represent mean±SEM. Taurine (14–126 mg/kg, i.p.) or diazepam (0.3–0.6 mg/kg, i.p.) was injected 30 min prior to the measurement of exploratory behavior.
⁎Pb0.05,⁎⁎Pb0.01 vs. vehicle-treated group (one-way ANOVA followed by two-tailed Dunnett' t-test). n=10–11.
273W.X. Kong et al. / Pharmacology, Biochemistry and Behavior 83 (2006) 271–276
and diazepam have no significant effect on the locomotor
activity in HU and HF conditions.
4.2. Hole-board test
Data are summarized in Table 1. ANOVA (df=5,55) showed
significant effects on number [F=3.58, Pb0.01] and duration
[F=2.80, Pb0.05] of head-dipping, number of rearing
[F=3.60, Pb0.01], rearing duration [F=2.55, Pb0.05] and
number of squares crossed [F=5.04, Pb0.01]. Compared with
the control group, further analyses confirmed that taurine
(42 mg/kg) significantly increased number (Pb0.01) and
duration (Pb0.05) of head-dipping. Rearing counts (Pb0.05)
and the number of squares crossed (Pb0.01) were also
significantly increased. Diazepam, at both 0.3 and 0.6 mg/kg,
significantly increased the number of squares crossed (Pb0.05
and Pb0.01). Analysis also revealed that administration of
0.6 mg/kg diazepam significantly increased number (Pb0.01)
and duration (Pb0.05) of rearing.
4.3. Open field test
The results for the open field test are shown in Fig. 2.
ANOVA demonstrated significant treatment effects on number
of center entries [F (4,44)=6.57, Pb0.01], time spent in central
area [F (4,43)=3.47, Pb0.05], and anti-thigmotactic effect
score [F (4,43)=5.25, Pb0.01]. Further analyses showed that
taurine (126 mg/kg) and diazepam (2 mg/kg) significantly
increased the number of center entries (both Pb0.01) and time
spent in central area (Pb0.01 and Pb0.05 respectively). See
Fig. 2A and B. Taurine (126 mg/kg) also significantly increased
the anti-thigmotactic score (Pb0.01) (See Fig. 2D). Fig. 2C
presents the total locomotor activity of vehicle-treated and drug-
treated rats exposed to a 5-min session in the open field test.
None of the drugs tested showed a significant effect on this
In the present study, taurine, at dose of 42 mg/kg, had
apparently anxiolytic properties in rats exposed to social inter-
action and hole-board tests. In the open field test, the effective
dose of taurine was higher (126 mg/kg) than the other two tests.
These observations suggested taurine produced several clear
anxiolytic effects in the battery of anxiety tests, although dif-
ferent tests were associated with some different outcomes.
The social interaction test was developed 27 years ago (File
and Hyde, 1978) as the first animal test of anxiety that used a
natural form of behavior as the dependent measure. An increase
in social interaction, without a concomitant increase in motor
activity, is indicative of an anxiolytic effect. Both benzodiaze-
pines and drugs acting on the 5-HT system have been found to
have effects in this test (Dunn et al., 1989). The dorsal hip-
pocampus played an important role in controlling behavior in
social interaction test. As previously stated, this animal model
provides the opportunity of varying the level of anxiety that
is generated by the test conditions. There is evidence for
increasing endogenous serotonergic tone but decreasing endog-
enous cholinergic tone in the dorsal hippocampus with in-
creasing anxiety. Thus, the dorsal hippocampal serotonergic and
Number of Central Entries
Time spent in Central Area(s)
Fig. 2. The effect of taurine (14, 42, 126 mg/kg) and diazepam (2.0 mg/kg) on rats behaviorin the open-field test: (A) the number of center entries; (B) time spent in the
central area of the open field; (C) motor activity; (D) anti-thigmotactic effect. All drugs were injected 30 min before behavior test. Results are expressed as means±
SEM. Significant differences from corresponding vehicle:⁎Pb0.05,⁎⁎Pb0.01. n=8–10. Veh: □, Taurine: ▨, Diazepam: ■.
274 W.X. Kong et al. / Pharmacology, Biochemistry and Behavior 83 (2006) 271–276
cholinergic systems are both biochemically and behaviorally
tightly coupled, and appear to have an antagonistic relationship
in the modulation of anxiety (File et al., 2000). Smythe et al.
(1996) reported that stimulation of benzodiazepine receptors
may offset the loss of cholinergic systems. In our present study,
taurine (42 mg/kg, i.p.) and diazepam (2 mg/kg, i.p.)
significantly increased the time of social interaction without
an increase in locomotion in HU condition, the highest level of
anxiety, indicating an anxiolytic effect. Based on other's studies
that exogenous taurine (10− 4M) inhibits release and synthesis
of newly formed serotonin 5-HT from tryptophan (Becquet et
al., 1993) and affects the binding of ligands to the benzodiaz-
epine site (Medina and De Robertis, 1984; Malminen and
Kontro, 1986), we presumed that the anxiolytic effect of taurine
in social interaction test may be due to decrease endogenous
serotonin tone on the one hand and increase endogenous choline
tone on the other hand while the anxiolytic effect of diazepam
was mainly concerned with the increasing endogenous choline
in dorsal hippocampus.
Takeda et al. (1998)indicatedthathead-dippingbehaviorwas
sensitive to changes in the emotional state of the animal, and
suggested that the expression of an anxiolytic state in animals
may be reflected by an increase in head-dipping behavior. In the
duration of head-dipping, the number of rearing, the number of
squares crossed in the hole-board test. It could be argued that the
increase of head-dipping in rats is merely an artifact of the
hyperactivity induced by the drug. In order to make clear the
facts,weexamined theeffect of taurine on thelocomotoractivity
of mice in a separate experiment. The results in mice showed
taurine had no influence on their locomotor activity at the
concurrently with anxiety (i.e., in the same test), sometimes the
measure used is not the most appropriate because it is a biased
measure, loading on both the “activity” and “anxiety” factors. In
our current study, diazepam, a putative anxiolytic agent,
as well as taurine. Therefore the increase of number of squares
crossed and rearing is not only a representation of hyperactivity
but also a reflection of anxiolytic effect.
During open field exposure, rats treated with 126 mg/kg
squares entered was not significantly affected by taurine. The
kg or more of taurine decreased ambulation and increased
thigmotaxis. The probable reason for this discrepancy is that the
previous study tested taurine in a high light illumination field
while the present study used a dim light arena. And the animals
effects of many drugs are dependent upon the environmental
conditions under which the test is conducted and the age of test
animals(Higgins et al., 1992; Serrano et al., 2002).
Additionally, there are some obvious differences on the
outcomes of the three anxiety tests, such as the locomotor
activity of the tested animals and the anxiolytic doses of TA.
Although various experimental models of anxiety (e.g. plus-
maze, social interaction, Vogel conflict, light/dark exploration,
hole-board, free-exploration, and neophobia tests) have been
proposed to measure different types or states of anxiety, there is
some uncertainty as to whether anxiety mechanisms and
anxiolytic drugs are uniformly active within and between
animal models (Handley and MaBlane, 1993). They often yield
variable or contradictory effects, probably as a result of
differences in the target receptor or subtypes, animal models,
dose range and routes of administration (Griebel, 1995).
In conclusion, the findings of the present study provide more
experimental evidences to suggest taurine has anxiolytic-like
effects on anxiety animal models. And this effect may be
mediated by the interaction of taurine with 5-HT and GABA
system, although this conclusion remains speculative in the
absence of neurochemical data. Taurine may act as a modulating
or anti-anxiety agent in the central nervous system but further
studies investigating the mechanism underlying the behavioral
actions of taurine may be necessary.
Baskin SI, Hinkamp DL, Marquis WJ, Tilson HA. Effects of taurine on
psychomotor activity in the rat. Neuropharmacology 1974;13:591–4.
Becquet D, Hery M, Francois-Bellan AM, Giraud P, Deprez P, Faudon M, et al.
Glutamate, GABA, glycine and taurine modulate serotonin synthesis and
release in rostral and caudal rhombencephalic raphe cells in primary
cultures. Neurochem Int 1993;23(3):269–83 [Sep].
Benrabh H, Bourre J-M, Lefauconnier J-M. Taurine transport at the blood-brain
barrier: an in vivo brain perfusion study. Brain Res 1995;692:57–65.
Bockelmann R, Reiser M, Wolf G. Potassium-stimulated taurine release and
nitric oxidesynthaseactivity duringquinolinic acidlesion of the rat striatum.
Neurochem Res 1998;23(4):469–75 [Apr].
Cheeta S, Kenny PJ, File SE. Hippocampal and septal injections of nicotine and
8-OH-DPAT distinguish among different animal tests of anxiety. Prog
Neuropsychopharmacol Biol Psychiatry 2000;24(7):1053–67 [Oct].
Chen SW, Kong WX, Zhang YJ, Li YL, Mi XJ, Mu XS. Possible anxiolytic
ColivicchiMA, BianchiL,Bolam JP, Galeffi F, Frosini M, Palmi M, et al. The in
vivo release of taurine in the striatonigral pathway. Adv Exp Med Biol
Darstein M, Landwehrmeyer GB, Kling C, Becker C-M, Feuerstein TJ.
Strychnine-sensitive glycine receptors in rat caudaputamen are expressed by
cholinergic interneurons. Neuroscience 2000;96:33–9.
Dunn RW, Corbett R, Fielding S. Effects of 5-HT1A receptor agonists and
NMDA receptor antagonists in the social interaction test and the elevated
plus maze. Eur J Pharmacol 1989;169:1-10.
File SE, Hyde JRG. Can social interaction be used to measure anxiety? Br
J Pharmacol 1978;62:19–24.
File SE, Kenny PJ, Cheeta S. The role of the dorsal hippocampal serotonergic
and cholinergic system in the modulation of anxiety. Pharmacol Biochem
Foos TM, Wu J-Y. The role of taurine in the central nervous system and the
modulation of intracellular calcium homeostasis. Neurochem Res
Galarreta M, Bustamante J, Martin del Rio R, Solis JM. Taurine induces a long-
lasting increase of synaptic efficacy and axon excitability in the hip-
pocampus. J Neurosci 1996;16(1):92-102 [Jan].
Griebel G. 5-Hydroxytryptamine-interacting drugs in animal models of anxiety
disorders: more than 30 years research. Pharmacol Ther 1995;65:319–95.
Handley SL, MaBlane JW. 5-HT drugs in animal models of anxiety.
275W.X. Kong et al. / Pharmacology, Biochemistry and Behavior 83 (2006) 271–276
Higgins GA, Jones BJ, Oakley NR. Effect of 5-HT1A receptor agonists in two Download full-text
models of anxiety after dorsal raphe injection. Psychopharmacology
Hruska RE, Thut PD, Huxtable RJ, Bressler R. Suppression of conditional
drinking by taurine and related compounds. Pharmacol Biochem Behav
Jacobsen JG, Smith LH. Biochemistry and physiology of taurine and taurine
derivatives. Physiol Rev 1968;48(2):424–511.
Kamei J, Ohsawa M, Tsuji M, Takeda H, Matsumiya T. Modification of the
effects of benzodiazepines on the exploratory behaviors of mice on a hole-
board by diabetes. Jpn J Pharmacol 2001;86:47–54.
Kontro P, Oja SS. Interactions of taurine with GABAB binding sites in mouse
brain. Neuropharmacology 1990;29(3):243–7 [Mar].
Kuriyama K. Symposium, taurine as a neuromodulator. Fed Proc 1980;39
Liljequist R. Interaction of taurine and related compounds with GABAergic
neurones in the nucleus raphe dorsalis. Pharmacol Biochem Behav
Lombardini JB. High affinity uptake systems for taurine in tissue slices and
synaptosomal fractions prepared from various regions of the rat central
nervous system. Correction of transport data by different experimental
procedures. J Neurochem 1977;29(2):305–12 [Aug].
Malminen O, Kontro P. Actions of taurine on the GABA-benzodiazepine
receptor complex by taurine in rat brain membranes. Neurochem Res
McBride WJ, Frederickson RC. Taurine as a possible inhibitory transmitter in
the cerebellum. Fed Proc 1979;39:2701–5.
Medina JH, De Robertis E. Taurine modulation of the benzodiazepine gamma-
aminobutyric acid receptor complex in brain membranes. J Neurochem
Michel HB, Richard WO. Taurine acts on a subclass of GABAAreceptors in
mammalian brain in vitro. Eur J Pharmacol 1991;207:9-16.
Milakofsky L, Hare AT, Miller MJ, Vogel HW. Rat plasma levels of amino acids
and released compounds during stress. Life Sci 1984;36:753–61.
Min L, Chen SW, Li WJ, Wang R, Li YL, Wang WJ, et al. The effects of
angelica essential oil in social interaction and hole-board tests. Pharmacol
Biochem Behav 2005;81(4):838–42 [Aug].
stratum measured by microdialysis. Amino Acids 2004;27(3–4):261–8 [Dec].
Oja SS, Kontro P. GABA, hypotaurine and taurine transport in brain slices from
developing mouse. Dev Neurosci 1984;6:271–7.
Sanberg PR, Ossenkopp K-P. Dose-response effects of taurine on some open
field behaviors in the rat. Psychopharmacology 1977;53:207–9.
Saransaari P, Oja SS. Characterization of N-methyl-D-aspartate-evoked taurine
release in the developing and adult mouse hippocampus. Amino Acids
Sergeeva OA, Haas HL. Expression and function of glycine receptors in striatal
cholineric interneurons from rat and mouse. Neuroscience 2001;104:1043–55.
Serrano MI, Goicoechea C, Serrano JS, Serrano-Martino MC, Sánchez E,
Martín MI. Age-related changes in the antinociception induced by taurine in
mice. Pharmacol Biochem Behav 2002;73:863–7.
Sgaragli G, Carla V, Magnani M, Galli A. Hypothermia induced in rabbits by
intracerebroventricular taurine: specificity and relationship with central
serotonin (5-HT) systems. J Pharmacol Exp Ther 1981;219:778–85.
SiemiątkowskiM, Sienkiewicz-jarosz H, CzłonkowskaAI,Bidziński A, Płaźnik
A. Effects of buspirone, diazepam, and zolpidem on open field behavior, and
brain [3H] muscimol binding after buspirone pretreatment. Pharmacol
Biochem Behav 2000;66(3):645–51.
Smythe JW, Murphy D, Costall B. Benzodiazepine receptor stimulation blocks
scopolamine-induced learning impairments in a water maze task. Brain Res
TakedaH, Tsuji M, Matsumiya T. Changes in head-dipping behaviorin the hole-
board test reflect the anxiogenic and/or anxiolytic state in mice. Eur J
276W.X. Kong et al. / Pharmacology, Biochemistry and Behavior 83 (2006) 271–276