The Journal of Neuroscience, 2001, Vol.

21 RC165 1 of 6

Neural Correlates of Conscious Self-Regulation of Emotion

Mario Beauregard,1,2,3 Johanne Lévesque,3,4 and Pierre Bourgouin1

1Département de Radiologie, Faculté de Médecine, Université de Montréal and Centre Hospitalier de l’Université de
Montréal (CHUM), Hôpital Notre-Dame, Montréal, Québec, Canada H2L 4M1, 2Centre de Recherche en Sciences
Neurologiques, Faculté de Médecine, Université de Montréal, Montréal, Québec, Canada H3C 3J7, 3Centre de
Recherche, Institut Universitaire de Gériatrie de Montréal, Montréal, Québec, Canada H3W 1W5, and 4Département de
Psychologie, Université de Montréal, Montréal, Québec, Canada H3C 3J7

A fundamental question about the relationship between cogni- ciated with activation of the right superior frontal gyrus and right
tion and emotion concerns the neural substrate underlying anterior cingulate gyrus. No activation was found in limbic
emotional self-regulation. To address this issue, brain activation areas. These findings reinforce the view that emotional self-
was measured in normal male subjects while they either re- regulation is normally implemented by a neural circuit compris-
sponded in a normal manner to erotic film excerpts or volun- ing various prefrontal regions and subcortical limbic structures.
tarily attempted to inhibit the sexual arousal induced by viewing They also suggest that humans have the capacity to influence
erotic stimuli. Results demonstrated that the sexual arousal the electrochemical dynamics of their brains, by voluntarily
experienced, in response to the erotic film excerpts, was asso- changing the nature of the mind processes unfolding in the
ciated with activation in “limbic” and paralimbic structures, psychological space.
such as the right amygdala, right anterior temporal pole, and Key words: emotional self-regulation; consciousness; voli-
hypothalamus. In addition, the attempted inhibition of the sex- tion; metacognition; prefrontal cortex; “limbic” structures; func-
ual arousal generated by viewing the erotic stimuli was asso- tional magnetic resonance imaging

Emotional self-regulation refers to the heterogeneous set of pro- the conduct of human affairs in daily life, few things are known
cesses by which individuals influence, consciously and voluntarily, with respect to the neurobiological substrate subserving the psy-
which emotions they have, when they have them, and how they chological processes associated with the volitional modulation of
experience and express these emotions (Gross, 1999). Given the emotional responses. With respect to this question, Nauta (1971)
multicomponential nature of emotions—which comprise cogni- and Tucker et al. (1995) have theorized that the frontal neocortex
tive, experiential, behavioral, and physiological dimensions— and limbic/paralimbic structures are components of a frontolim-
emotional self-regulation may be associated with changes in one bic network involved in the regulation of emotion. More recently,
or more of these response domains (Gross, 1999). This form of Davidson and associates (2000) have proposed, on the basis of
self-regulation involves decreasing, maintaining, or increasing various evidence from experimental lesion studies in animals and
both negative and positive emotions (Masters, 1991; Parrott, clinical neuropsychological, psychophysiological, and functional
1993; Langston, 1994), by using various cognitive regulatory pro- brain mapping studies in humans, that emotional self-regulation
cesses such as rationalization, reappraisal, and suppression may be normally implemented by a neural circuit consisting of
(Gross, 1999; Hariri et al., 2000). several regions of the prefrontal cortex (PFC) [e.g., orbitofrontal
Disturbances of normal emotional self-regulation might be a PFC, dorsolateral PFC, anterior cingulate cortex (ACC)], and of
key factor in the genesis of depression and anxiety, both of which subcortical limbic structures, such as the amygdala and the hypo-
may involve a chronic inability to suppress negative emotion thalamus. The present functional magnetic resonance imaging
(Jackson et al., 2000). Regarding this issue, it has been postulated (fMRI) study was undertaken to test the validity of this view.
that impulsive aggression and violence also arise as a conse-
MATERIALS AND METHODS
quence of defective regulation of emotional responses (Davidson
Subjects. Ten healthy male (C aucasian) volunteers (age range, 20 – 42;
et al., 2000). Given that the ability to modulate emotions is at the mean age, 23.5) participated in this study. Informed consent was obtained
heart of the human experience, it is not surprising that the use of from all subjects.
emotional self-regulatory processes constitutes the core of several
modern psychotherapeutic approaches.
Despite the immense impact of emotional self-regulation for This article is published in The Journal of Neuroscience, Rapid
Communications Section, which publishes brief, peer-
Received March 8, 2001; revised June 4, 2001; accepted June 28, 2001. reviewed papers online, not in print. Rapid Communications
This work was supported by grants from National Sciences and Engineering are posted online approximately one month earlier than they
Research Council of Canada (NSERC) and Département de Radiologie, Faculté de
Médecine, Université de Montréal to M.B. We thank the staff of the Département
would appear if printed. They are listed in the Table of
de Radiologie, Centre Hospitalier de l’Université de Montréal (CHUM), Hôpital Contents of the next open issue of JNeurosci. Cite this article
Notre-Dame, for their skillful technical assistance. as: JNeurosci, 2001, 21:RC165 (1–6). The publication date is
Correspondence should be addressed to Dr. Mario Beauregard, Centre de Re-
cherche, Institut Universitaire de Gériatrie de Montréal, 4565 Queen Mary Road, the date of posting online at www.jneurosci.org.
Montréal, Québec, Canada, H3W lW5. E-mail: beauregm@magellan.umontreal.ca.
Copyright © 2001 Society for Neuroscience 0270-6474/01/210001-06$15.00/0 http://www.jneurosci.org/cgi/content/full/5619
2 of 6 J. Neurosci., 2001, Vol. 21 Beauregard et al. • Neural Correlates of Emotion Regulation

Behavioral procedures. Blood oxygen level-dependent (BOLD) signal space with a three-dimensional isotropic Gaussian kernel (12 mm f ull
changes were measured during two experimental conditions, i.e., a sexual width half maximum) to improve the signal-to-noise ratio and to accom-
arousal condition and an attempted inhibition condition. In the sexual modate for residual variations in f unctional neuroanatomy that usually
arousal condition, subjects viewed a series of erotic film excerpts and persist between subjects after spatial normalization.
emotionally neutral film excerpts. They were instructed to react normally For both sexual arousal and attempted inhibition conditions, a con-
to both types of stimuli. In the case of the erotic film excerpts, this meant ventional subtraction method was used to contrast the brain activity
that the subjects had to allow themselves to become sexually aroused in associated with the viewing of the erotic film excerpts and that associated
response to the erotic film excerpts. In the attempted inhibition condi- with the viewing of the emotionally neutral film excerpts. For the
tion, subjects were instructed to inhibit any emotional reaction to both statistical analysis, the time series of the images were correlated with the
categories of stimuli. These instructions meant that the subjects had to delayed boxcar f unction that approximates the activation patterns, and a
voluntarily decrease the intensity of the sexual arousal felt in reaction to fixed effects general linear model for autocorrelated observations was
the erotic film excerpts. To accomplish that goal, subjects were encour- applied voxelwise. Voxel values for each contrast yielded a statistical
aged to distance themselves from these stimuli, that is, to become a parametric map of the t statistic (SPM t), subsequently transformed to the
detached observer. They were also instructed to look at the stimuli unit normal distribution (SPM Z). An a priori search strategy was used.
directly during both experimental conditions. A whole-brain post hoc analysis was also performed. The a priori search
Erotic film excerpts were selected on the basis of previous work by our strategy predicted significant activation in the PFC (orbitofrontal PFC,
group demonstrating the efficiency of such stimuli at bringing rapidly and dorsolateral PFC, anterior cingulate cortex), the amygdala, the temporal
automatically a marked positive change in the subjective emotional pole, and the hypothalamus. The search volume corresponding to the
experience of healthy male subjects, change correlated with significant brain regions of interest was defined a priori by tracing the neuroana-
activation of limbic brain regions (Beauregard et al., 1998). These film tomical boundaries of these regions on the MR reference image. These
excerpts depicted different kinds of sexual interactions (e.g., oral sex, boundaries were also traced using the standard atlas of Talairach and
vaginal intercourse, etc.) between one woman and two or three men, two Tournoux (1988). For this a priori search, a corrected probability thresh-
women and one man, and between two or more women. The emotionally old of p ⬍ 0.05 was used. For the whole-brain post hoc analysis, a
neutral film excerpts were matched to the erotic film excerpts with corrected probability threshold of p ⬍ 0.005 was used. Only clusters
respect to the number and the gender of the individuals involved in showing a spatial extent of at least 10 contiguous voxels were kept for
various social interactions (e.g., interviews, carpentry, etc). Erotic film image analysis.
excerpts were presented with their original sound tracks consisting of
instrumental music. To avoid any confounding effect attributable to the
information processing of language perceived in an auditory manner, RESULTS
emotionally neutral music was superimposed on the neutral film excerpts.
Both categories of film excerpts were selected from a private bank of
Behavioral data
validated stimuli created by the investigators. Phenomenologically, the viewing of the erotic film excerpts in
The sexual arousal condition corresponded to one f unctional run and both the sexual arousal and attempted inhibition conditions in-
consisted of the viewing of, first, four blocks of emotionally neutral film duced a transient state of sexual arousal in all subjects. The mean
excerpts and, then, four blocks of erotic film excerpts. On the basis of
evidence gathered previously by our group, this design was adopted to level of reported sexual arousal was significantly higher in the
avoid contamination of the neutral stimuli by the erotic stimuli. Each sexual arousal condition (mean ⫽ 5; range, 3– 8) than in the
block lasted 39 sec. Blocks were separated by resting periods of 15 sec, inhibition condition (mean ⫽ 2; range, 1– 4) ( p ⬍ 0.005). For both
during which subjects viewed a blank cyan screen. The attempted inhi- experimental conditions, the viewing of the erotic film excerpts
bition condition corresponded to the other f unctional run and also
did not produce changes in the emotional state other than sexual
consisted of the viewing of, first, four blocks of emotionally neutral film
excerpts and, then, four blocks of erotic film excerpts. The order of arousal. Likewise, in both of these conditions, the viewing of the
presentation of the f unctional runs was counterbalanced across subjects. emotionally neutral film excerpts did not generate primary emo-
In addition, the neutral and erotic stimuli that were used in the sexual tional responses. Furthermore, the strategy questionnaire indi-
arousal condition for half of the subjects were presented for the other cated that, in the attempted inhibition condition, most subjects
half of the subjects in the attempted inhibition condition (and vice versa).
To assess the subjective responses of the subjects to the stimuli, succeeded at distancing themselves from the erotic stimuli, i.e., at
immediately at the end of each f unctional run, they were asked to rate becoming a detached observer.
verbally — on a visual analog rating scale ranging from 0 (absence of any
emotional reaction) to 8 (strongest emotion ever felt in one’s lifetime)— fMRI data
the average intensity of sexual arousal or primary emotions [e.g., sadness,
happiness, disgust, fear, anger, surprise (Plutchik, 1994)] felt during the A priori search
viewing of both categories of film excerpts. For each f unctional run, an When the brain activity associated with the viewing of the emo-
average rating score was computed for the four blocks of erotic film tionally neutral film excerpts was subtracted from that associated
excerpts and the four blocks of emotionally neutral film excerpts. At the
with the viewing of the erotic film clips in the sexual arousal
end of the scanning session, subjects were also asked to complete a
“strategy questionnaire” in which they described the emotion regulation condition, significant BOLD signal increases were found in the
strategies they used to inhibit the sexual arousal generated by the erotic right amygdala, the right anterior temporal pole [Brodmann area
stimuli. (BA) 38], and the hypothalamus (Fig. 1, Table 1). In addition,
MR I. Echoplanar images (EPIs) were acquired on a 1.5 Tesla system when the brain activity associated with the viewing of the emo-
(Magnetom Vision, Siemens Electric, Erlangen, Germany). T wenty-
eight slices (5 mm thick) were acquired every 3 sec in an inclined axial tionally neutral film excerpts was subtracted from that associated
plane, aligned with the anterior commissure –posterior commissure axis. with the viewing of the erotic film clips in the attempted inhibi-
These T2* weighted f unctional images were acquired using an EPI pulse tion condition, significant loci of activation were noted in the right
sequence [repetition time (TR) ⫽ 0.8 msec; echo time (TE) ⫽ 54 msec; superior frontal gyrus (BA 10) and the right anterior cingulate
Flip angle ⫽ 90°; field of view (FOV) ⫽ 215 mm; Matrix ⫽ 64 ⫻ 64)].
gyrus (BA 32) (Fig. 2, Table 1). No significant loci of activation
After functional scanning, high-resolution data were acquired via a Tl-
weighted three-dimensional volume acquisition obtained using a gradient were detected in the amygdalas, the anterior temporal polar
echo pulse sequence (TR ⫽ 9.7 msec; TE ⫽ 4 msec; Flip angle ⫽ 12°; region, and the hypothalamus (even for a low height threshold
FOV ⫽ 215 mm; Matrix ⫽ 256 ⫻ 256). value of p ⬍ 0.1, uncorrected). With regard to individual data, the
Data were analyzed using Statistical Parametric Mapping software same pattern was found; that is, right prefrontal activation was
(SPM99, Wellcome Department of Cognitive Neurology, L ondon, UK).
Images for all subjects were realigned to correct for artifacts caused by
seen in all 10 subjects, whereas no activation was noted in the
small head movements and spatially normalized into an MRI stereotactic amygdala and the hypothalamus, and activation in the anterior
space (Talairach and Tournoux, 1988). Images were then convolved in temporal pole was noted in only one subject.
Beauregard et al. • Neural Correlates of Emotion Regulation J. Neurosci., 2001, Vol. 21 3 of 6

Figure 1. Statistical activation maps

are shown for limbic–paralimbic struc-
tures defined a priori. Images are coro-
nal sections for the data averaged across
subjects. The right hemisphere of the
brain corresponds to the right side of the
image. In the Sexual Arousal condition,
greater activation during the viewing of
erotic film excerpts relative to the view-
ing of emotionally neutral film excerpts
was noted in the right amygdala ( A),
right anterior temporal pole ( B), and
hypothalamus ( C). In the Attempted In-
hibition condition, no significant loci of
activation were seen in the amygdalas
( D), anterior temporal polar region ( E),
and hypothalamus ( F).

Table 1. BOLD signal increases in the search volume defined a priori (p < 0.05, corrected)

Talairach coordinates (mm)

Region Brodmann area x y z Z statistic
Sexual arousal condition R amygdala 25 ⫺3 ⫺14 4.17
R anterior temporal pole 38 32 20 ⫺26 4.05
Hypothalamus 1 ⫺7 ⫺9 3.97
Attempted inhibition condition R superior prefrontal gyrus 10 22 46 13 4.25
R anterior cingulate gyrus 32 9 42 4 3.8

Stereotaxic coordinates are derived from the human atlas of Talairach and Tournoux (1988) and refer to medial –lateral position (x) relative to medline (positive ⫽ right),
anterior–posterior position (y) relative to the anterior commissure (positive ⫽ anterior), and superior–inferior position (z) relative to the commissural line (positive ⫽
superior). Designation of Brodmann areas for cortical areas are also based on this atlas. L, Left; R, right.

Whole-brain post hoc analysis DISCUSSION

When the brain activity associated with the viewing of the emo- In agreement with the results of a previous study by our group
(Karama et al., 2001), the viewing of erotic film excerpts, in the
tionally neutral film excerpts was subtracted from that associated
sexual arousal condition, produced a significant activation of the
with the viewing of the erotic film clips in the sexual arousal
right amygdala, right anterior temporal pole (BA 38), and hypo-
condition, significant BOLD signal increases were seen in the left
thalamus. These findings fit with the various lines of evidence
middle occipital gyrus (BA 19), the right superior occipital gyrus
suggesting that the amygdala plays a pivotal role in the evaluation
(BA 19), the right inferior temporal gyrus (BA 37), the left of the emotional significance of stimuli (for review, see Lane and
cerebellum, and the superior parietal lobule (BA 7), bilaterally Nadel, 2000), that the hypothalamus is a key brain structure
(Table 2). Moreover, when the brain activity associated with the implicated in the neuroendocrine and autonomic expression of
viewing of the emotionally neutral film excerpts was subtracted emotion (Carter, 1998), and that the anterior temporal pole is
from that associated with the viewing of the erotic film clips in the involved in the imparting of emotional color to subjective expe-
attempted inhibition condition, significant loci of activation were rience (Mesulam, 1985). In light of the anatomic projections that
found in the left middle occipital gyrus (BA 19), the right inferior the central nucleus of the amygdala sends to the hypothalamus,
and superior occipital gyri (BA 19), the left superior occipital and the basolateral nuclei of the amygdala sends to the temporal
gyrus (BA 19), the left inferior frontal gyrus (BA 44), and the pole (Martin, 1996), it appears conceivable that in the sexual
superior parietal lobule (BA 7), bilaterally (Table 2). arousal condition, the right amygdala, the hypothalamus, and the
4 of 6 J. Neurosci., 2001, Vol. 21 Beauregard et al. • Neural Correlates of Emotion Regulation

Figure 2. Statistical activation maps showing peaks of acti-

vation in prefrontal cortical regions defined a priori during the
Attempted Inhibition condition. Images are coronal sections
for the data averaged across subjects. The right hemisphere of
the brain corresponds to the right side of the image. Signifi-
cant loci of activation were noted in the right superior frontal
gyrus ( A) and right anterior cingulate gyrus ( B).

Table 2. BOLD signal increases in the whole-brain post hoc analysis ( p < 0.005, corrected)

Talairach coordinates (mm)

Brodmann
Region area x y z Z statistic
Sexual arousal condition L middle occipital gyrus 19 ⫺48 ⫺72 0 19.2
R inferior temporal gyrus 37 51 ⫺66 ⫺9 17.48
R superior occipital gyrus 19 30 ⫺70 33 11.9
R superior parietal lobule 7 27 ⫺57 60 11.85
L superior parietal lobule 7 ⫺30 ⫺54 60 11.83
L cerebellum ⫺24 ⫺84 21 11.28
Attempted inhibition condition L middle occipital gyrus 19 ⫺48 ⫺72 ⫺3 13.71
R inferior occipital gyrus 19 45 ⫺72 3 11.01
R superior occipital gyrus 19 30 ⫺78 36 10.11
L superior parietal lobule 7 ⫺27 ⫺54 54 9.49
L superior occipital gyrus 19 ⫺21 ⫺84 21 9.38
R superior parietal lobule 7 30 ⫺54 60 7.68
L inferior frontal gyrus 44 ⫺57 9 30 5.67

L, Left; R, right.

right anterior temporal pole are functional components of a for a low height threshold value of p ⬍ 0.1, uncorrected). Fur-
neural circuit subserving the processing of the erotic visual stim- thermore, the analysis of individual data revealed the same pat-
uli. Within such a network, we surmise that, first, the right tern, i.e., no significant loci of activation was found in the amyg-
amygdala participated in the evaluation of the emotional content dala, the anterior temporal polar region, and the hypothalamus
of the complex perceptual information associated with the visual whenever the right superior frontal gyrus and right anterior
processing of the erotic film excerpts. Second, this brain structure cingulate gyrus were activated. These findings provide some
sent a message to the hypothalamus. Such modulation of the support for the hypothesis that the dorsolateral PFC, the amyg-
hypothalamic activity then led to sexual arousal (Sachs and dala, and the hypothalamus participate in a neural circuit in-
Meisel, 1984). Third, for this physiological arousal to become volved in emotional self-regulation (Davidson et al., 2000). They
fully conscious, in addition to sending outputs to the hypothala- are also consistent with the results of a recent functional brain
mus, the amygdala sent, in parallel, a message to the right anterior imaging study showing that a cognitive task, such as choosing
temporal pole. Such a role for this brain region concurs with labels that identify the emotional expression of faces, correlated
recent work showing that this paralimbic associative cortical area with simultaneous increased regional cerebral blood flow (rCBF)
is correlated with attending to one’s own emotional experience in the right prefrontal cortex and decreased rCBF in the amyg-
(Lane, 2000). dalas (Hariri et al., 2000). In other respects, these results are
The results of the attempted inhibition condition revealed that consistent with findings indicating that the dorsolateral PFC is
the volitional inhibitory action produced by the male subjects, to implicated in metacognitive/executive top-down processes, which
decrease the intensity of the sexual arousal produced by the erotic refer to the ability to monitor and control the information pro-
film excerpts, was associated with activation peaks in BA 10 of the cessing necessary to produce voluntary action (Flavell, 1979).
right dorsolateral PFC (superior frontal gyrus) and BA 32 of These results also accord with the view that, by virtue of its
the right ACC. Contrary to the pattern of brain activity seen in anatomic connections with brain regions implicated in the mod-
the sexual arousal condition, in the attempted inhibition condi- ulation of autonomic and endocrine functions, such as the amyg-
tion, no significant loci of activation were noted in the amygdalas, dala, hypothalamus, and orbitofrontal PFC, the rostral-ventral
the anterior temporal polar region, and the hypothalamus (even affective subdivision of the ACC is primarily involved in the
Beauregard et al. • Neural Correlates of Emotion Regulation J. Neurosci., 2001, Vol. 21 5 of 6

evaluation of emotional and motivational information and in the dorsolateral PFC, right ACC, right amygdala, right anterior tem-
regulation of emotional responses (Vogt et al., 1992; Devinsky et poral pole, and hypothalamus may constitute a fundamental
al., 1995; Bush et al., 2000). psychobiological mechanism through which human beings can
The whole-brain post hoc analysis revealed that in addition to consciously and willfully self-regulate their emotional responses,
producing activation in the right amygdala, right anterior tempo- using various metacognitive processes. From a phylogenetic per-
ral pole, and hypothalamus, the sexual arousal condition also spective, such a circuit may implement one of the most remark-
induced significant activation in the inferotemporal cortex, the able human faculties that has emerged in the course of human
extrastriate visual cortex, the cerebellum, and the superior pari- evolution. At both an individual and a collective level, a defect of
etal lobule. Moreover, in addition to producing activation in the this neural circuitry (or of an analogous network involving a
right superior frontal gyrus and the right anterior cingulate gyrus, prefrontal cortical modulation of negative emotional responses
the attempted inhibition condition was also associated with acti- associated with activation in limbic structures) may have disas-
vation in the extrastriate visual cortex, the superior parietal trous psychological and social consequences. Ontologically, the
lobule, and the left inferior frontal gyrus. present findings suggest that humans have the capacity to influ-
Clinical neuropsychological data have demonstrated that cere- ence the electrochemical dynamics of their brains, by voluntarily
bellar lesions can produce a blunting of effect (Schmahmann and changing the nature of the mind processes unfolding in the
Sherman, 1998). In light of these data, and on the basis of the fact psychological space.
that cerebellar activation has often been noted in functional brain Last, we acknowledge some of the limitations that characterize
imaging studies of emotion generated by emotionally laden visual the present study. First, the restriction to one arousal state makes
stimuli, one could argue that the cerebellar activation found here it difficult to argue for the specificity, or generality, of the brain
suggests that the cerebellum is somehow related to the “feeling” findings for both the sexual arousal and attempted inhibition
experience associated with sexual arousal. Further studies are conditions. Second, because there was no experimental condition
awaited to tackle this intriguing question. in which subjects had to voluntarily increase their arousal state, it
With respect to the inferotemporal cortex, the results of a remains possible that the activations seen in the attempted inhi-
recent magnetoencephalography study (Streit et al., 1999) suggest bition condition reflected an effort to modulate the sexual arousal
that this cortical region is involved in the recognition of the induced by the erotic stimuli rather than an actual inhibition of
various facial expressions of emotions. Given these findings and this arousal.
the nature of the erotic film excerpts used in this study, it seems
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