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The Brain Functional Networks Associated to Human and
Animal Suffering Differ among Omnivores, Vegetarians
and Vegans
Massimo Filippi1,2*, Gianna Riccitelli1, Andrea Falini3, Francesco Di Salle4, Patrik Vuilleumier5, Giancarlo
Comi2, Maria A. Rocca1,2
1 Neuroimaging Research Unit, Institute of Experimental Neurology, Division of Neuroscience, Scientific Institute and University Hospital San Raffaele, Milan, Italy,
2 Department of Neurology, Scientific Institute and University Hospital San Raffaele, Milan, Italy, 3 Department of Neuroradiology, Scientific Institute and University
Hospital San Raffaele, Milan, Italy, 4 Maastricht Brain Imaging Center, Department of Cognitive Neuroscience, University of Maastricht, Maastricht, The Netherlands,
5 University Medical Center of Geneva, University of Geneva, Geneva, Switzerland
Abstract
Empathy and affective appraisals for conspecifics are among the hallmarks of social interaction. Using functional MRI, we
hypothesized that vegetarians and vegans, who made their feeding choice for ethical reasons, might show brain responses
to conditions of suffering involving humans or animals different from omnivores. We recruited 20 omnivore subjects, 19
vegetarians, and 21 vegans. The groups were matched for sex and age. Brain activation was investigated using fMRI and an
event-related design during observation of negative affective pictures of human beings and animals (showing mutilations,
murdered people, human/animal threat, tortures, wounds, etc.). Participants saw negative-valence scenes related to
humans and animals, alternating with natural landscapes. During human negative valence scenes, compared with
omnivores, vegetarians and vegans had an increased recruitment of the anterior cingulate cortex (ACC) and inferior frontal
gyrus (IFG). More critically, during animal negative valence scenes, they had decreased amygdala activation and increased
activation of the lingual gyri, the left cuneus, the posterior cingulate cortex and several areas mainly located in the frontal
lobes, including the ACC, the IFG and the middle frontal gyrus. Nonetheless, also substantial differences between
vegetarians and vegans have been found responding to negative scenes. Vegetarians showed a selective recruitment of the
right inferior parietal lobule during human negative scenes, and a prevailing activation of the ACC during animal negative
scenes. Conversely, during animal negative scenes an increased activation of the inferior prefrontal cortex was observed in
vegans. These results suggest that empathy toward non conspecifics has different neural representation among individuals
with different feeding habits, perhaps reflecting different motivational factors and beliefs.
Citation: Filippi M, Riccitelli G, Falini A, Di Salle F, Vuilleumier P, et al. (2010) The Brain Functional Networks Associated to Human and Animal Suffering Differ
among Omnivores, Vegetarians and Vegans. PLoS ONE 5(5): e10847. doi:10.1371/journal.pone.0010847
Editor: Pedro Antonio Valdes-Sosa, Cuban Neuroscience Center, Cuba
Received March 19, 2010; Accepted May 5, 2010; Published May 26, 2010
Copyright: ß 2010 Filippi et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: The authors have no support or funding to report.
Competing Interests: The authors have declared that no competing interests exist.
* E-mail: filippi.massimo@hsr.it
suggests that the merging between emotions and feelings
experienced by oneself and those perceived in other individuals
may be a key ingredient of social understanding, and it may play
a major role in promoting empathy, prosocial behaviours, and
moral norms [1,3]. Moreover, empathic responses can be
modulated by the subjective attitude held toward suffering
individuals [7], as well as by personal experience [8]. Several
functional magnetic resonance imaging (fMRI) studies showed
that observing the emotional state of another individual activates
a neuronal network involved in processing the same state in
oneself, whether it is pain, disgust, or touch[3,4,5]. Empathy
toward another person, which can be defined as the ability to
share the other person’s feeling in an embodied manner, has been
related to recruitment of a network mostly including the
somatosensory and insular cortices, limbic regions and the
anterior cingulate cortex (ACC). Whereas cognitively inferring
about the state of other person (known as theory of mind) has
been associated with recruitment of medial prefrontal regions, the
superior temporal sulcus and the temporo-parietal junction[4].
Introduction
Social cognition includes mental processes necessary to
understand and store information about the self and other
persons, as well as interpersonal norms and procedures to
navigate efficiently in the social world [1]. Basic abilities
underlying social cognition include the perception and evaluation
of social stimuli, the integration of perceptions with contextual
knowledge, and finally the representation of possible responses to
the situation. One of the hallmarks of social cognition in humans
is the ability to understand conspecifics as beings like oneself, with
intentional and mental lives like one’s own [2]. Accordingly,
human beings tend to identify with conspecifics and attribute
mental states to them. Such abilities rely on the activity of several
brain regions, including the frontal lobes (orbitofrontal cortex,
medial prefrontal cortex, and cingulate cortex), the temporal
lobes (including the amygdala), the fusiform gyrus, and the
somatosensory cortices [3,4,5]. The majority of these regions is
also critically involved in the processing of emotions [6]. This
PLoS ONE | www.plosone.org
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May 2010 | Volume 5 | Issue 5 | e10847
fMRI and Feeding Habits
A few investigations have also assessed whether affective links
between people modulate their brain empathic responses to others,
such as when these are loved ones or strangers[9], or when they
are believed to be fair or unfair persons [7,9]. The majority of
previous studies attempting to characterize empathy-related
responses did not separate empathy towards humans from that
towards animals. Furthermore, in some studies, scenes showing
animals were treated as a neutral condition. However, a recent
study [10] that compared stimuli depicting human and non
human animal targets demonstrated higher subjective empathy as
the stimuli became closer in phylogenetic relatedness to humans
(mammalian vs. bird stimuli), thus indicating that empathic
response towards humans may generalize to other species.
In this study, we postulated that the neural representation of
conditions of abuse and suffering might be different among
subjects who made different feeding choice due to ethical reasons,
and thus result in the engagement of different components of the
brain networks associated with empathy and social cognition. In
details, we tested the hypothesis that the neural processes
underlying empathy in vegetarians and vegans may not only
operate for representations about humans but also animals, and
thus vary between them and omnivore subjects. Vegetarians and
vegans, who decided to avoid the use of animal products for
ethical reasons, have a moral philosophy of life based on a set of
basic values and attitudes toward life, nature, and society, that
extends well beyond food choice. The earliest records of
vegetarianism as a concept and practice among a significant
number of people was closely connected with the idea of
nonviolence towards animals and was promoted by religious
groups and philosophers. The term veganism, which was coined
from vegetarianism, acknowledges the intrinsic legitimacy of all
sentient life and rejects any hierarchy of acceptable suffering
among creatures. Veganism is a lifestyle that seeks to exclude the
use of animals for food, clothing, or any other purpose [11]. The
central ethical question related to veganism is whether it is right
for humans to use and kill animals. Due to these differences of
believes and behaviours, we also hypothesized that, in addition to
a common shared pattern of cortical processing of human and
animal suffering, vegetarians and vegans might also have
functional architecture differences reflecting their different
motivational factors and believes.
Figure 1. Graph showing error bars of means and standard
deviations of empathy quotient (EQ) score in the three groups
of subjects. See text for further details.
doi:10.1371/journal.pone.0010847.g001
Between-group fMRI results
The patterns of activations during the neutral condition did not
differ between groups.
Common regions of activations between vegetarians and
vegans
During human negative valence picture view, omnivore
subjects had a more significant activation (p,0.05, FWE) of the
bilateral middle temporal gyrus (MTG) (MNI space coordinates:
38, 258, 8, t value = 5.65; and 236, 276, 8, t value = 5.56) when
compared to vegetarians and vegans. Compared to omnivore
subjects, the entire sample of vegetarians and vegans had more
significant activations (p,0.05, FWE) of the ACC (MNI space
coordinates: 10, 22, 40; 10, 36, 28, and 24, 30, 36; t
values = 5.65, 5.43, and 5.30), and the left inferior frontal gyrus
(IFG) (MNI space coordinates: 248, 20, 0, t value = 5.56)
(Figure 3).
During animal negative valence picture view, omnivore subjects
had more significant activations (p,0.05, FWE) of the bilateral
MTG (MNI space coordinates: 246, 262, 0, t value = 6.03; and
34, 274, 4, t value = 5.94), when compared to vegetarians and
vegans. Compared to omnivore subjects, the entire sample of
vegetarians and vegans had more significant activations (p,0.05,
FWE) of the bilateral IFG (MNI space coordinates: 250, 14, 22, t
value = 6.84; and 52, 14, 24, t value = 6.34), bilateral lingual gyrus
(MNI space coordinates: 8, 280, 214, t value = 6.83; and 210,
278, 214, t value = 6.58), ACC (MNI space coordinates: 0, 24,
28; 22, 52, 8; t values = 5.76 and 5.51), posterior cingulate cortex
(PCC) (MNI space coordinates: 0, 242, 26, t value = 5.87), left
cuneus (MNI space coordinates: 22, 278, 24, t value = 5.83), and
left middle frontal gyrus (MFG) (MNI space: 244, 46, 8, t
value = 5.50) (Figure 3). This analysis also showed that, compared
to omnivores, vegetarians and vegans had a lower activation of the
right amygdala (MNI space coordinates: 30, 2, 220, t
value = 5.38). To better define amygdala behavior in the three
groups of subjects, we analyzed its activations and deactivations
during the two experimental conditions in each group (Tables 1
and 2). This analysis revealed no significant activation neither
deactivation (even when lowering the threshold for the statistical
Results
Empathy assessment
The Empathy quotient (EQ) score was significantly different
between groups (p = 0.002). At post-hoc analysis, the EQ score was
significantly higher in vegetarians in comparison with omnivore
subjects (mean EQ score = 49.5, SD = 8.9 in vegetarians vs. 38.8,
SD = 8.1 in omnivore; p = 0.001), and in vegans (mean EQ
score = 44.6, SD = 9.8) in comparison with omnivore subjects
(p = 0.04) (Figure 1). The difference between vegans and
vegetarians was not statistically significant.
Within-group fMRI results
The observation of both human and animal negative valence
scenes resulted in the recruitment of several brain areas involved in
emotion and empathy in the three groups of subjects, including the
anterior insula, basal ganglia, thalami, and several other cortical
areas located in the occipital lobes, prefrontal and parietal cortices.
Figure 2 shows the brain patterns of activations in the three groups
of subjects during the different experimental conditions. Table 1
summarizes the main results of within-group comparisons of the
two experimental conditions.
PLoS ONE | www.plosone.org
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May 2010 | Volume 5 | Issue 5 | e10847
fMRI and Feeding Habits
Figure 2. Within-group analysis of activations. Cortical activations on a rendered brain from omnivore (A–H), vegetarian (I–R) and vegan (S–W)
subjects during observation of pictures showing negative valence scenes of humans (A–D, I–N, S–V) or animals (E–H, O–R, Z–W) (within-group
analysis, one-sample t tests, t = 3 for display purpose). Images are in neurological convention.
doi:10.1371/journal.pone.0010847.g002
significance at a p,0.001, uncorrected) during animal picture
view in this region in vegetarians and vegans.
Analysis of interaction
To further explore the specificity of stimulus processing within the
three groups of subjects, we performed an analysis of interaction
between picture types (animal/human) and groups (omnivore/
vegetarian/vegan). Results showed an interaction in the right
amygdala (MNI space coordinates: 24, 210, 222) (greater increases
to animal negative valence view in omnivores and to human negative
valence view in vegans) (Figure 3), the left amygdala (222, 28, 228)
(greater increases to human negative valence view in vegans)
(Figure 4), the ACC (MNI space coordinates: 22, 52, 10) (preferential
increases to human negative valence view in omnivores, and to
animal negative valence view in vegetarians) (Figure 4); and the right
IFG (MNI space coordinates: 52, 20, 28) (selective responses to
animal negative valence view in vegans) (Figure 4).
Table 2 summarizes the behavior, in terms of activations/
deactivations, at the within-group one sample t test analysis of the
three main areas which showed a significant interaction between
groups and conditions (i.e., amygdala, IFG, and ACC).
Different regions of activations between vegetarians and
vegans
We also directly compared the neural responses in empathy and
emotion-related networks between omnivores, vegetarians, and
vegans, using a masking procedure (See Methods), to identify
regions of specific activations of each group contrasted to the
others.
a) Vegetarians vs. omnivores and vegans. Observation of
human negative valence scenes resulted in a selective recruitment
of the right IPL (BA40) (MNI space coordinates: 52, 250, 40, t
value = 4.44) in vegetarians (Figure 3). For animal pictures,
activations specific to vegetarians were found in the ACC (MNI
space coordinates: 22, 52, 10, t value = 5.02) and the right lingual
gyrus (MNI space coordinates: 8, 284, 210, t value = 5.00)
(p,0.05, FWE).
b) Vegans vs. omnivores and vegetarians. During human
negative valence picture view, no cortical activation ‘‘specific’’ to
vegans was found. During animal negative valence picture view,
vegans activated the IFG bilaterally (MNI space coordinates: 54,
16, 26, and 246, 18, 22, t values = 4.88 and 4.67), and the left
MFG (BA10) (MNI space coordinates: 246, 48, 4, t value = 4.29)
(Figure 3) (p,0.05, FWE).
PLoS ONE | www.plosone.org
Analysis of correlations
During human negative valence picture view, no correlation
was found between EQ score and fMRI activity in the three
groups of subjects of the study.
During animal negative picture view, significant correlations
(p,0.001) were found between EQ score and:
3
May 2010 | Volume 5 | Issue 5 | e10847
fMRI and Feeding Habits
Table 1. Within-group comparisons of human vs. animal negative valence picture view and vice versa in omnivore subjects,
vegetarians and vegans (paired t test in each group, p,0.05 FWE-corrected).
Human vs. animal pictures
Animal vs. human pictures
Omnivore
Vegetarians
Vegans
Omnivore
Vegetarians
Vegans
MNI
coordinates
X Y
Z
MNI
coordinates
X
Y
Z
MNI
coordinates
X
Y
Z
MNI
coordinates
X
Y
Z
MNI
coordinates
X
Y
Z
Activation sites
BA
MNI
coordinates
X
Y
Z
R amygdala


R MTG
37
46,
L MTG
37
248, 274,
264,
30, 216, 222
26,
28, 218
22,


2
60, 232, 2
44,
56,
264, 10
8,
228



26

262, 250, 2
252, 22, 222



24,
226
R lingual gyrus
19
14,
254,
22





L lingual gyrus
19
210, 266,
22





R cuneus
18
16,
280,
28
14, 286,
R precuneus
7
8,
254,
43

R insula


38, 24,
R thalamus



28
8








20,
226, 0
26, 24
R putamen



30,
L putamen



228, 6,
R IOG




210













250, 266, 214
20,
286,
212
L IOG




230, 282,
210

L IPL
40



248, 256,
46


R MFG




38,
50


4,
L MFG




240, 14,
36


R IFG




44,
28

46,
ACC
32




0,
50,
PCC
23




2,
250, 20
44,
10
22,
28
240,
26
0,
MNI = Montreal Neurological Institute, R = right, L = left, BA = Brodmann area, MTG = middle temporal gyrus, IOG = inferior occipital gyrus, IPL = inferior parietal lobule,
MFG = middle frontal gyrus, IFG = inferior frontal gyrus, ACC = anterior cingulate cortex, PCC = posterior cingulate cortex.
doi:10.1371/journal.pone.0010847.t001
N
N
N
activation of the left MTG (r = 0.87), ACC (r = 20.76) and the
bilateral IFG (right IFG: r = 20.71, left IFG: r = 20.89) in
omnivores;
activation of the left IFG (r = 0.92), the left MFG (r = 0.68),
and the right MTG (r = 20.75) in vegetarians;
activation of the bilateral lingual gyrus (right lingual gyrus:
r = 0.69, left lingual gyrus: r = 0.75) and the left IFG (r = 0.78)
in vegans.
regarding animals rather than humans, with the additional
recruitment of the mPFC, PCC, and some visual areas. ACC
has been associated with alert states, self awareness and pain
processing [12], whereas mPFC and PCC activations are
frequently observed in conditions involving representation of the
self and self values [13]. The PCC is also thought to be involved in
memory and visuospatial processing [14], particularly in relation
to emotions and social behavior [13]. PCC is consistently activated
when subjects have to judge the valence of emotionally salient
words or episodic memories, with the strongest responses seen
when unpleasant stimuli are presented [14].
The notion that empathic response might differ among
vegetarians, vegans and omnivores, and that such a response
might vary during viewing of human and animal sufferance is at
least partially supported by the results of EQ assessment in the
three groups of subjects and by the analysis of correlation between
EQ scores and fMRI findings, which showed a direct relationship
between the EQ score and left IFG recruitment during animal
suffering view in vegetarians and vegans, whereas in omnivores
such a relationship was inverse.
The pattern of increased recruitment of empathy-related areas
in vegetarians and vegans during animal suffering view was also
associated with a reduced activation of the right amygdala in
comparison to omnivores. The amygdala responds to various kinds
of aversive stimuli, most strongly fearful and threatening scenes
[15] and, to a lesser extent, to those associated with disgust [16].
Discussion
The first main finding of this study was the demonstration of a
common functional architecture of emotional processing in
vegetarians and vegans. In particular, while omnivores are
characterized by a greater activation of the bilateral posterior
MTG during both human and animal negative valence scenes,
vegetarians and vegans have constantly an higher engagement of
empathy related areas while observing negative scenes, independently of the species of the individuals involved, which is
characterized by an increased recruitment of the ACC and the
IFG. Increased activation in the ACC and left IFG in vegetarians
and vegans during human and animal suffering view is likely to
reflect a stronger empathic response in the first two groups.
Remarkably, vegetarians and vegans have an higher engagement of empathy related areas while observing negative scenes
PLoS ONE | www.plosone.org
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May 2010 | Volume 5 | Issue 5 | e10847
fMRI and Feeding Habits
Figure 3. Results of the between-group compa …
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