Psychoneuroendocrinology (2012) 37, 87—93

a v a i l a b l e a t w w w. s c i e n c e d i r e c t . c o m

j o u r n a l h o m e p a g e : w w w. e l s e v i e r. c o m / l o c a t e / p s y n e u e n

Oxytocin specifically enhances valence-dependent

parasympathetic responses
Matthias Gamer *, Christian Büchel

Department of Systems Neuroscience, University Medical Center Hamburg-Eppendorf, Martinistr. 52, D-20246 Hamburg, Germany

Received 11 January 2011; received in revised form 11 April 2011; accepted 9 May 2011

KEYWORDS Summary The evolutionarily highly conserved neuropeptide oxytocin seems to be involved in
Oxytocin; the regulation of complex forms of social behavior. It enhances the processing of positive social
Facial expression; stimuli, reduces behavioral and neuroendocrine stress responses and modulates amygdala
Emotion; activity in humans. Moreover, it has been proposed that oxytocin dampens sympathetic nervous
Heart rate; system activity. This hypothesis was tested in a double-blind, placebo-controlled study with 38
Skin conductance; men either receiving 24 IU oxytocin intranasally or a placebo spray. While accomplishing an
Autonomic nervous system emotion classification task, electrodermal responses were measured as an index of sympathetic
activity. Moreover, heart rate changes were recorded that are additionally mediated by the
parasympathetic nervous system. Oxytocin enhanced differential heart rate responses to facial
expressions as a function of the emotional valence, but had no effect on electrodermal activity or
tonic measures of physiological arousal. These results indicate that oxytocin specifically mod-
ulates phasic activity of the parasympathetic nervous system which potentially reflects an
increased motivational value of facial expressions following oxytocin treatment. Findings suggest
that anxiolytic effects of oxytocin are not reflected in short-term sympathetic responses and may
even be a consequence of rather than a prerequisite for improved social information processing.
# 2011 Elsevier Ltd. All rights reserved.

phobia, psychopathy, autism; cf., Clark and McManus, 2002;

1. Introduction
Social relations are an integral part of human society and
promote mental health. Impairments in social perception
and behavior play a crucial role in the etiology and main-
tenance of a number of psychiatric disorders (e.g., social
* Corresponding author at: Department of Systems Neuroscience,
University Medical Center Hamburg-Eppendorf, Bldg. W34, Marti-
nistr. 52, D-20246 Hamburg, Germany. Tel.: +49 40 7410 57160;
fax: +49 40 7410 59955.
E-mail address: m.gamer@uke.uni-hamburg.de (M. Gamer).
Brüne and Brüne-Cohrs, 2006). Because of this key role,
research on the neurobiological substrates of these pro-
cesses has intensified in recent years (Adolphs, 2010). The
evolutionarily highly conserved neuropeptide oxytocin has
been identified as one important biological factor regulat-
ing complex forms of social behavior (Insel, 2010). Oxytocin
reduces anxiety and neuroendocrine stress responses and it
facilitates approach behavior across several species (Bartz
and Hollander, 2006; Heinrichs and Gaab, 2007). It mod-
ulates memory for social information and enhances the
processing of social cues (Heinrichs et al., 2009). These
diverse psychosocial functions seem to be at least in part
mediated by the amygdala (e.g., Kirsch et al., 2005; Domes

0306-4530/$ — see front matter # 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.psyneuen.2011.05.007
88
et al., 2007a; Gamer et al., 2010; Labuschagne et al.,
2010).
The anxiolytic properties of the neuropeptide as well as its
influence on amygdala activation may also imply that oxyto-
cin modulates the sympathetic or parasympathetic tone. In
fact, oxytocin was found to decrease subjective arousal
ratings for threatening social stimuli (Norman et al., 2010)
and it reduces the functional connectivity between the
amygdala and the brain stem regions implicated in autonomic
manifestations of emotional experience (Kirsch et al., 2005).
Based on these observations, it seems likely that oxytocin
primarily reduces activity of the sympathetic nervous system
which closely reflects the arousal state of the organism
(Dawson et al., 2007). To our knowledge, however, this
question has not been empirically tested until now. In the
present double-blind placebo-controlled study, we continu-
ously measured skin conductance responses and brief heart
rate changes while participants classified emotional facial
expressions. This setup allowed us to simultaneously examine
oxytocin effects on both branches of the autonomic nervous
system because skin conductance is exclusively modulated by
the sympathetic nervous system (Wallin, 1981) whereas heart
rate changes are additionally mediated by the parasympa-
thetic nervous system (Berntson et al., 2007). We expected
that oxytocin should reduce sympathetically mediated skin
conductance responses and this effect might be more pro-
nounced to fearful facial expression (cf., Norman et al.,
2010). An explorative analysis of phasic heart rate responses
was conducted to test whether oxytocin differentially affects
both branches of the autonomic nervous system. Since pre-
vious studies showed that oxytocin specifically enhances
attention to the eyes of conspecifics (Domes et al., 2007b;
Guastella et al., 2008; Gamer et al., 2010), we additionally
manipulated whether participants initially fixated on the eye
or mouth region. Thereby, we were able to determine
whether the autonomic nervous system responds differen-
tially to specific facial features in the focus of attention.
2. Methods
2.1. Participants and pharmacological
manipulation
Forty-six right-handed, male volunteers participated volun-
tarily in this study that was approved by the ethics committee
of the medical faculty of the University of Rostock. Within
this study, functional magnetic resonance imaging data and
eye movements were recorded in addition to autonomic
data. The current study focuses specifically on modulatory
effects of oxytocin on autonomic nervous system activity and
the results pertaining neuroimaging data and eye movements
are reported elsewhere (Gamer et al., 2010). For the current
analysis, eight subjects were excluded because of severe
recording artifacts that did not allow for a reliable quanti-
fication of autonomic responses. The final sample consisted
of 38 men (mean age  SD = 24.6  3.5 years). All subjects
were healthy and did not report a history of neurological or
endocrine disease, medication, and drug or alcohol abuse.
They were instructed to abstain from alcohol and nicotine for
12 h before the experiment and they were paid for participa-
tion. Participants in the oxytocin and the placebo group did
M. Gamer, C. Büchel
not differ with respect to age, education or occupation (see
online supplement).
The pharmacological manipulation was realized in a dou-
ble-blind between-subjects design (N = 19 in each group)
with participants either receiving a single intranasal dose
of 24 IU oxytocin (Syntocinon spray; Novartis; three puffs per
nostril, each with 4 IU oxytocin) or a placebo spray that
contained all inactive ingredients except for the neuropep-
tide. The experimental task that lasted no longer than 45 min
started approximately 45 min after substance administra-
tion. State questionnaires were used to control for nonspe-
cific effects of oxytocin (see Supplementary Materials and
Methods). Similar to previous studies (e.g., Domes et al.,
2007a,b; Guastella et al., 2008), oxytocin did neither mod-
ulate positive or negative effect nor wakefulness, arousal or
mood (Figure S1).
2.2. Experimental task
The experimental task was based on a fully crossed 3  2
design with the factors emotional expression and initial
fixation. Using published validation ratings, we selected male
and female faces unambiguously depicting neutral, fearful
and happy expressions from four established data sets: Nim-
Stim Face Stimulus Set (www.macbrain.org), Pictures of
Facial Affect (Ekman and Friesen, 1971), Karolinska Directed
Emotional Faces (Lundqvist et al., 1998), and FACES database
(Ebner et al., 2010). These faces were slightly rotated such
that both eyes had exactly the same height in each image.
Colored images were converted to grayscales and an elliptic
mask was fitted to solely reveal the face itself while hiding
hair and ears. Finally, the cumulative brightness was normal-
ized across images.
A different sample of 72 individual faces (12 males and 12
females for each expression) was randomly selected from this
pool of pictures for each participant. These faces were then
presented once in a randomized sequence in each of three
experimental blocks. Each block lasted approximately
11 min. The number of female and male faces within each
block was similar for the different emotional expressions. A
trial started with a fixation cross (2 s) followed by the pre-
sentation of the face (150 ms) followed by a blank screen
(1850 ms) and another fixation cross for a randomly chosen
period of 2—9 s. To precisely control for the initial fixation,
half of the male and female stimuli within each emotional
expression were unpredictably shifted either downward or
upward on each trial such that the eyes or the mouth
appeared at the location of the fixation cross (Gamer and
Büchel, 2009; Gamer et al., 2010). The presentation duration
of facial expressions was considerably shorter than in pre-
vious studies (e.g., Kirsch et al., 2005; Domes et al., 2007a)
to ensure that the initially fixated feature remained in the
focus of attention during picture presentation. Any saccade
that is triggered by the facial expression would occur after
stimulus offset. Participants were instructed to classify the
depicted emotional expression as quickly and accurately as
possible by pressing the corresponding key on a button box
with the right hand. The keys were labeled according to the
emotional expression. Participants were instructed to press
the index, middle and ring finger when recognizing fearful,
neutral, or happy expressions, respectively. Although stimuli
 Oxytocin modulates parasympathetic activity
Figure 1 Peripheral physiological responses to emotional expressions in the placebo and the oxytocin group. (A) Phasic heart rate changes for 8 poststimulus seconds. (B) Average heart
rate changes and (C) log-transformed skin conductance response amplitudes as a function of the emotional expression and the initial fixation. Error bars indicate SEM.

89
90
were only shown very briefly, participants performed at
ceiling with no influence of oxytocin on classification accu-
racy or response times (see online supplement for a detailed
analysis of the behavioral data).
2.3. Data acquisition and response scoring
During the emotion classification task, heart rate and skin
conductance were measured continuously. These measures
were selected because they reflect activity of both branches
of the autonomic nervous system and they can be easily
acquired within an MRI setting where the current experiment
was conducted. Although we were mainly interested in phasic
responses of the autonomic nervous system, tonic measures
were also derived from a 10 s baseline period preceding each
experimental block. Exploratory analyses were conducted to
test whether oxytocin modulates the general arousal level.
However, the currently derived tonic measures should only be
regarded as an approximation of tonic autonomic activity and
future studies should include a separate baseline period for
measuring tonic activity of the sympathetic and parasympa-
thetic nervous system.
Heart rate (HR) recording was accomplished by means of
an electrocardiogram (ECG). The electrodes were placed
according to the recommendations of the manufacturer of
the physiologic ECG measurement unit (Siemens) left of the
sternum approximately at the level of the 3rd intercostal
space as well as on the lower rib cage left and right of the
midclavicula line. This setup allowed for simultaneously
measuring ECG lead I and aVF. ECG data were recorded at
400 Hz and saved for offline analysis. To quantify phasic
stimulus related HR changes, R-waves were detected from
the average of both leads and R—R intervals were converted
to HR (in beats per minute, bpm). Afterwards a real time
scaling procedure was applied (Velden and Wölk, 1987)
resulting in one HR value for each 500 ms segment of a period
from 0 to 8 s following stimulus onset. The HR in the last
second prior to stimulus onset represented the prestimulus
baseline and poststimulus difference scores (DHR) were
derived by subtracting the prestimulus baseline value from
the HR-score of each poststimulus value. In addition to phasic
HR changes, we also estimated tonic HR by calculating the
mean HR during a period of 10 s preceding stimulus presenta-
tion in each experimental block. These values were averaged
within each participant.
Skin conductance was measured by a constant voltage
system (0.5 V) using a bipolar recording with two Ag/AgCl
electrodes placed on the thenar and hypothenar surfaces of
the participant’s left hand. The electrodermal data was
initially digitized with 100 Hz but subsequently down-
sampled to 10 Hz for offline analysis. To obtain phasic sti-
mulus related skin conductance responses (SCRs), we
decomposed the recording into a set of SCRs using an indi-
vidually fitted response template for each participant (Lim
et al., 1997). The procedure resulted in a set of SCRs for each
electrodermal recording that best resembled the measured
data. Subsequently, SCRs that were elicited by the stimuli
were identified by searching for responses with a latency of 1
and 3 s after stimulus onset. The amplitudes of these
responses were log-transformed to account for the positive
skew of the distribution (Venables and Christie, 1980). To
M. Gamer, C. Büchel
examine potential effects of oxytocin on tonic electroder-
mal activity, the skin conductance recording was low pass
filtered using a cutoff frequency of 0.05 Hz. Subsequently,
the mean skin conductance level was calculated in a period
of 10 s preceding stimulus presentation in each experimen-
tal block. Corresponding to the heart rate analysis, these
values were averaged within each participant.
Trials with excessive artifacts that impeded a detection of
R-waves or a quantification of SCR amplitudes were excluded
from the analysis. On average, autonomic responses could be
analyzed for 97% of all trials (SD = 1%). Blood oxygenation
level dependent functional images and eye movements were
additionally acquired during the task but these data are
reported elsewhere (Gamer et al., 2010).
2.4. Statistical analyses
The phasic heart rate responses were analyzed using a
2  3  2  16 analysis of variance (ANOVA) incorporating
the between-subject factor drug (oxytocin vs. placebo)
and the within-subject factors emotional expression (fearful,
happy, neutral), initial fixation (eyes vs. mouth) and time
window (500 ms segments in a period from 0 to 8 s following
stimulus onset). Amplitudes of skin conductance responses
were analyzed using a 2  3  2 ANOVA with the between-
subject factor drug (oxytocin vs. placebo) and the within-
subject factors emotional expression (fearful, happy, neu-
tral) and initial fixation (eyes vs. mouth). Tonic heart rate and
skin conductance level were compared between the oxytocin
and the placebo group by means of t-tests for independent
samples.
In ANOVAs involving more than one degree of freedom in
the enumerator, the Huynh—Feldt procedure was applied to
correct for potential violations of the sphericity assumption.
Cohen’s f or Cohen’s d are reported as effect size estimates.
3. Results
As can be seen from Fig. 1A, HR changes showed a biphasic
pattern consisting of an initial increase that passed into a
deceleration approximately 3—4 s after stimulus onset. The
ANOVA revealed significant main effects of emotional expres-
sion (F[2,72] = 6.70, e = 1.00, p = .008, f = 0.09), and time
window (F[15,540] = 45.37, e = .15, p < .001, f = 0.61).
Importantly, we additionally obtained a significant drug by
emotional expression interaction (F[2,72] = 3.20, e = 1.00,
p = .047, f = 0.06) and a marginally significant main effect
of drug (F[1,36] = 3.65, p = .064, f = 0.16) indicating that
overall heart rate responses were slightly larger in the
oxytocin group. All other effects did not reach statistical
significance (all p > .159, all f < 0.10).
To further clarify the drug by emotional expression inter-
action, post hoc oneway ANOVAs using emotional expression
as within-subject factor were carried out for each treatment
group separately. As depicted in Fig. 1B, mean HR changes
were comparable across experimental manipulations in the
placebo group and the ANOVA did not reveal a significant
effect of emotional expression (F[2,36] = 0.54, e = .97,
p = .582, f = 0.06). In the oxytocin group, however, HR
changes were modulated by the emotional expression and
the ANOVA revealed a significant effect (F[2,36] = 9.80,
Oxytocin modulates parasympathetic activity
e = 1.00, p < .001, f = 0.30). Fearful — and to a smaller
degree also neutral faces — resulted in overall HR accelera-
tions relative to happy faces (fearful vs. happy faces,
t[18] = 4.19, p < .001, d = 0.73; neutral vs. happy faces,
t[18] = 2.77, p = .013, d = 0.47). A marginally significant dif-
ference was also found between fearful and neutral facial
expressions (t[18] = 1.73, p = .099, d = 0.26). Interestingly,
such oxytocin dependent differentiation of physiological
responses as a function of emotional valence could not be
obtained for skin conductance response amplitudes and the
ANOVA did not reveal any statistically significant effect (all
p > .110, all f < 0.08, see Fig. 1C).
Oxytocin specifically modulated phasic stimulus related
activity of the cardiovascular system. Neither tonic HR
(oxytocin: M  SD = 62.65  8.96 bpm; placebo M  SD =
61.04  8.85 bpm; t[36] = 0.56, p = .580, d = 0.18), nor skin
conductance level differed between groups (oxytocin:
M  SD = 12.41  13.65 mS; placebo: M  SD = 13.47 
11.98 mS; t[36] = 0.26, p = .800, d = 0.08).
4. Discussion
This study examined the influence of oxytocin on autonomic
responses in an emotion classification task. In contrast to our
expectations, there was no effect of oxytocin on tonic elec-
trodermal activity or on phasic skin conductance responses.
Thus, oxytocin does not seem to modulate activity of the
sympathetic nervous system. By contrast, we observed a
specific effect of the neuropeptide on phasic heart rate
changes. Participants in the oxytocin group showed a slightly
larger overall heart rate increase to emotional expressions as
compared to the placebo group. More importantly, however,
oxytocin enhanced differential heart rate responses as a
function of the emotional valence of the depicted expres-
sion. This effect was independent of the initial fixation and it
emerged quickly within 1—2 s after stimulus onset. Such
rapid heart rate changes are mediated by the parasympa-
thetic nervous system that has a much wider and faster
control over cardiac chronotropy than the sympathetic ner-
vous system (Berntson et al., 2007). Thus, this differential
pattern of electrodermal and cardiovascular responses indi-
cates that oxytocin specifically modulates phasic activity of
the parasympathetic nervous system.
Since no effect of the initial fixation was found, these
results indicate that oxytocin primarily modulates cardiovas-
cular effects of emotional valence processing. Attentional
processes during face perception, that were also found to be
modulated by oxytocin on the behavioral and neural level
(e.g., Guastella et al., 2008; Gamer et al., 2010), do not
seem to be mirrored in autonomic responses. Moreover, we
did not observe oxytocin effects on tonic measures of auto-
nomic nervous system activity although global effects of
systemically administered oxytocin on cardiovascular
responses were reported previously in rats (e.g., Costa-E-
Sousa et al., 2005). It remains unclear whether these global
effects are induced by an oxytocin modulation of the central
nervous system (Petersson et al., 1999) or directly related to
oxytocin receptor binding at the heart (Favaretto et al.,
1997). The absence of corresponding effects in the current
study may be due to the intranasal application of low doses of
oxytocin that primarily modulate central oxytocin levels
91
instead of massively enhancing the peripheral availability
of the neuropeptide.
The specific modulation of valence-dependent parasympa-
thetic responses in the current study may indicate that oxyto-
cin facilitated assigning motivational value to facial
expressions Previous studies repeatedly observed that differ-
ential heart rate responses reflect the motivational value of
the stimulus material. Stronger parasympathetic influences on
cardiac chronotropy were thought to occur when a stimulus
captures attention and drives motivated approach or avoid-
ance behavior (Bradley et al., 2001). Relative heart rate
decelerations — as observed for happy compared to fearful
and neutral faces under oxytocin treatment — might thus
reflect the detection of salient and behaviorally relevant
stimuli in the environment. Since such valence dependent
modulation of parasympathetic responses was absent in the
placebo group, these results seem to indicate that oxytocin
facilitates a motivational differentiation between positive
social stimuli (happy faces) that prompt more elaborate eva-
luation and are capable of initiating approach tendencies and
negative stimuli (fearful faces) that become less relevant and
are potentially avoided. The lack of an effect of oxytocin on
overt recognition accuracy in the current study, which is
consistent with previous studies using relatively easy experi-
mental tasks (e.g., Labuschagne et al., 2010; Kirsch et al.,
2005), indicates that such influence of oxytocin is not neces-
sarily related to an increased ability of cognitively differen-
tiating between emotional expressions (cf., Hurlemann et al.,
2010). However, the hypothesized motivational effect might
still result in enhanced approach behavior as recently demon-
strated by an increased desire for future social engagement
following positive social contact under oxytocin treatment
(Alvares et al., 2010). Especially when social stimuli are more
difficult to be interpreted, oxytocin seems to additionally
improve the cognitive ability to infer the mental state of
others from subtle facial cues (Domes et al., 2007b). The
current results suggesting that oxytocin facilitates a motiva-
tional differentiation of emotional expressions are also well in
line with a recent study demonstrating that oxytocin is capable
of enhancing learning from social cues and attributed this
effect to a modulation of the ascribed motivational value
(Hurlemann et al., 2010). However, future studies are neces-
sary to directly test this assumption by involving a behavioral
measure of approach and avoidance behavior.
Our data indicate that anxiolytic properties of oxytocin
which were observed behaviorally and with respect to the
activity of the hypothalamic—pituitary—adrenal (HPA) axis
(e.g., Heinrichs et al., 2003) are not directly related to a
short-term dampening of sympathetic activation. Thus, it
seems possible that the anxiolytic effect of oxytocin in
humans might be the consequence instead of the prerequisite
of adequate social information processing. In line with this
reasoning, centrally released oxytocin might directly
enhance the processing of facial cues (Domes et al.,
2007b), potentially by increasing gaze toward the eye region
(Guastella et al., 2008; Gamer et al., 2010). This would allow
for assigning motivational value to social information and
reduce uncertainty in social situations. Anxiolytic effects
would then result from this optimized information processing
and be reflected in a reduction of medium-term physiological
stress responses such as cortisol release (Heinrichs et al.,
2003). It has to be mentioned, however, that the current
92
results are only based on a sample of male participants. Since
oxytocin seems to have differential effects on the neural
circuitry involved in face processing in women (Domes et al.,
2010), our data might not generalize to females.
Oxytocin has been successfully used as an experimental
therapeutic agent in patients with autism spectrum disorder.
It improved emotion recognition abilities in these patients
(Guastella et al., 2010), facilitated socially appropriate
behavior and increase gaze toward the eye region (Andari
et al., 2010). These results were interpreted as resulting
from anxiolytic effects of the neuropeptide which allowed for
a more comprehensive processing of social situations. How-
ever, it also seems possible that such behavioral effects of
oxytocin precede reductions of anxiety and discomfort. In
this case, oxytocin treatment might also be promising for
other mental disorders that are accompanied by dysfunc-
tional biases in the interpretation of social situations (e.g.
social anxiety disorder; Clark and McManus, 2002) even when
such situations are not generally experienced as aversive
(e.g. borderline personality disorder; Linehan, 1995). In this
case, oxytocin would not be used to induce a general ten-
dency to approach social situations. Instead, oxytocin might
help to improve the interpretation of social situations by
reducing dysfunctional biases with the aim to facilitate
adequate motivated approach or avoidance behavior. It
remains an interesting challenge for future studies to exam-
ine this possibility using online measures of physiological
arousal such as electrodermal or cardiovascular responses
during social information processing. Since women have a
higher prevalence for a number of clinical conditions where
oxytocin treatment might be helpful (e.g., borderline per-
sonality disorder), future studies should include both sexes
instead of specifically focusing on males.
Role of the funding source
The study was supported by a grant from the Bundesminis-
terium für Bildung und Forschung (BMBF, Network ‘‘Social
Cognition’’). Christian Büchel is additionally funded by the
German Research Foundation (DFG) SFB TRR 58, project B3.
These funding sources had no further role in study design; in
the collection, analysis and interpretation of data; in the
writing of the report; and in the decision to submit the paper
for publication.
Conflict of interest
The authors have no biomedical financial interests or poten-
tial conflicts of interest.
Acknowledgement
We thank Gina Gutsche for her help and support during data
analysis.
Appendix A. Supplementary data
Supplementary data associated with this article can be
found, in the online version, at doi:10.1016/j.psyneuen.
2011.05.007.
M. Gamer, C. Büchel
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