8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 1/12
This article was downloaded by: [186.104.215.92]On: 16 February 2013, At: 06:07Publisher: RoutledgeInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House37-41 Mortimer Street, London W1T 3JH, UK
Social NeurosciencePublication details, including instructions for authors and subscription information:
http://www.tandfonline.com/loi/psns20
Neural responses to perceiving suffering in humans
and animalsRobert G. Franklin Jr.
a , Anthony J. Nelson
b , Michelle Baker
b , Joseph E. Beeney
b ,
Theresa K. Vesciob , Aurora Lenz-Watson
b & Reginald B. Adams Jr.
b
a Department of Psychology, Brandeis University, Waltham, MA, USA
b Department of Psychology, The Pennsylvania State University, University Park, PA, USA
Version of record first published: 13 Feb 2013.
To cite this article: Robert G. Franklin Jr. , Anthony J. Nelson , Michelle Baker , Joseph E. Beeney , Theresa K. Vescio ,
Aurora Lenz-Watson & Reginald B. Adams Jr. (2013): Neural responses to perceiving suffering in humans and animals, SocialNeuroscience, DOI:10.1080/17470919.2013.763852
To link to this article: http://dx.doi.org/10.1080/17470919.2013.763852
PLEASE SCROLL DOWN FOR ARTICLE
Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions
This article may be used for research, teaching, and private study purposes. Any substantial or systematicreproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to
anyone is expressly forbidden.The publisher does not give any warranty express or implied or make any representation that the contentswill be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses shouldbe independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims,proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly inconnection with or arising out of the use of this material.
8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 2/12
SOCIAL NEUROSCIENCE, 2013http://dx.doi.org/10.1080/17470919.2013.763852
Neural responses to perceiving suffering in humansand animals
Robert G. Franklin Jr.1 , Anthony J. Nelson2 , Michelle Baker 2 , Joseph E. Beeney2 ,
Theresa K. Vescio2 , Aurora Lenz-Watson2 , and Reginald B. Adams Jr.2
1Department of Psychology, Brandeis University, Waltham, MA, USA2Department of Psychology, The Pennsylvania State University, University Park, PA, USA
The human ability to perceive and understand others’ suffering is critical to reinforcing and maintaining our social
bonds. What is not clear, however, is the extent to which this generalizes to nonhuman entities. Anecdotal evidence
indicates that people may engage in empathy-like processes when observing suffering nonhuman entities, but
psychological research suggests that we more readily empathize with those to whom we are closer and more
similar. In this research, we examined neural responses in participants while they were presented with pictures of
human versus dog suffering. We found that viewing human and animal suffering led to large overlapping regions
of activation previously implicated in empathic responding to suffering, including the anterior cingulate gyrus
and anterior insula. Direct comparisons of viewing human and animal suffering also revealed differences such
that human suffering yielded significantly greater medial prefrontal activation, consistent with high-level theory of
mind, whereas animal suffering yielded significantly greater parietal and inferior frontal activation, consistent with
more semantic evaluation and perceptual simulation.
Keywords: Empathy; Emotion; Mentalizing; Anthropomorphism.
Perceiving suffering in others can elicit powerfulresponses in observers. We readily empathize with
the suffering of others, feeling their pain as our own.
Anecdotal evidence suggests that this extends to ani-
mals as well, as images of animal suffering evoke
strong responses in humans. Stories of animal suffer-
ing and abuse capture media headlines as do stories
of human suffering. For example, in the midst of
the devastating earthquake and tsunami in Japan in
March 2011, a video of a dog staying with another
suffering dog captured as much international atten-
tion as any story of human suffering. Additionally,
animal rights groups often present explicit photos of
abused animals in an attempt to effect change in animal
treatment policies. Even fictitious images of animal
Correspondence should be addressed to: Robert G. Franklin, Jr., Department of Psychology, Brandeis University, 415 South Street, Mailstop
062, Waltham, MA 02454, USA. E-mail: [email protected] and Reginald B. Adams, Jr., Department of Psychology, 464 Moore Building,
University Park, PA 16802, USA. E-mail: [email protected]
This research was supported by a Social Science Research Institute grant, Penn State University, to R. B. A., Jr. We acknowledge Amanda
Gearhart and David Pennell for their help with data collection and Jasmine Boshyan for her helpful comments on an earlier version of this
article.
suffering can produce horror and outrage in viewers.Director Francis Ford Coppola said of The Godfather ,
“Thirty people were shot in the movie, but people
only talked about ‘cruelty to animals,’ ” which was a
response to the severed horse head found in a char-
acter’s bed (as quoted in Kohn, 1990). Anecdotal
evidence suggests that animal suffering arouses similar
responses as human suffering (e.g., Kennedy, 1992);
however, some psychological research indicates that
thinking about human mental states is distinct from
thinking about animal states (e.g., Caramazza &
Shelton, 1998). This points to an intriguing question;
namely, do images of suffering in humans and animals
elicit similar empathy-related processes at the neural
level?
© 2013 Taylor & Francis
8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 3/12
2 FRANKLIN ET AL.
SUFFERING IN HUMANS ANDANIMALS
A large social and political movement decries human-
caused animal suffering, calling for the end of the
use of animals for food, as beasts of burden, and forsport (Singer, 1975). This movement is largely based
on the philosophical and ethical premise that animal
suffering is morally equivalent to human suffering
(Singer, 1974). Despite anecdotal evidence and ethi-
cal appeals suggesting the equivalency of human and
animal suffering, research on how humans perceive
suffering has almost exclusively examined responses
to other humans. This is partly because humans are
assumed to have a set of mental experiences that are
distinct from those that animals possess (Kennedy,
1992). This is also partly due to the fact that we pro-
cess our knowledge of humans in different, and neu-
rally dissociable, cognitive substrates compared with
knowledge of nonhuman beings or inanimate objects
(Caramazza & Shelton, 1998). For instance, judging if
a word describes a possible action of a human agent
yields greater activation in medial prefrontal regions,
whereas judging if a word describes an action a dog
can perform leads to greater activation in more tem-
poral regions (Mason, Banfield, & Macrae, 2004).
This suggests that thinking about humans and thinking
about animals recruit qualitatively different processes.
In the context of this study, the aforementioned
findings suggest that processing suffering in humans
may lead to greater activation of networks involvedin understanding others’ mental states. Several mental
processes are necessary to understand others’ suffer-
ing. These include the ability to decode what another
is thinking, represent an individual’s mental and/or
emotional state, take that person’s perspective, reg-
ulate emotions generated through identification with
another’s suffering, and maintain awareness of the
boundaries between self and others (Decety, 2007).
These processes elicit activation in a variety of brain
regions, three of which are particularly relevant to the
present research.
First, viewing others in pain consistently activates
the dorsal anterior cingulate (dACC) and the ante-rior insula (AI). These regions are part of a network
involved in alerting organisms of threats and dan-
gers in their environment and representing one’s own
emotional states. Specifically, the anterior cingulate
and AI are part of a network involved with rep-
resentations of self and others’ affective states and
are important in homeostatic regulation (Craig, 2002;
Singer, Critchley, & Preuschoff, 2009). The dACC is
involved in error detection and performance monitor-
ing (Holroyd et al., 2004). In regard to perceiving
threats in the environment and empathy more specifi-
cally, the dACC is thought to be akin to a neural alarm
system, active when something is “off” physiologi-
cally (Eisenberger & Lieberman, 2004). In addition,
the dACC is involved in emotion regulation (Davidson,
Putnam, & Larson, 2000). The AI is involved withintegrating somatic and emotional information and is
active when witnessing others in pain, as well as being
active in situations evoking disgust and a sense of
unfairness (Ostrowsky et al., 2002). These regions are
consistently found in studies of empathy for pain and
are thought to reflect a core network involved with
perceiving the suffering of others, whether this knowl-
edge is based on actually perceiving someone in pain
or if a person is told that another is suffering (Lamm,
Decety, & Singer, 2011). Further, shared activation in
these regions supports the hypothesis that the vicarious
experience of pain involves some of the same neural
processes as directly experiencing pain (Singer et al.,2004).
Second, brain regions involved with decoding what
others are thinking are involved in empathy for the suf-
fering of others. The medial prefrontal cortex (mPFC)
is important to mentalizing, or understanding the
behaviors of oneself or others in terms of mental
states, and is active in many studies involving decod-
ing what another is feeling (Amodio & Frith, 2006).
More apropos to perceiving suffering in others, learn-
ing a close other is in pain leads to mPFC activation
(Singer et al., 2004), perspective taking, and empathy
arousal (Cheng, Chen, Lin, Chou, & Decety, 2010).
In regard to empathy, mPFC activation may reflect
the importance of self-awareness and taking the per-
spective of others, as lesions of the ventral mPFC
and orbitofrontal cortex lead to deficits in empathy
(Eslinger, 1998; Rankin et al., 2006; Sturm, Rosen,
Allison, Miller, & Levenson, 2006) and these regions
are involved in taking another’s emotional perspec-
tive (Hynes, Baird, & Grafton, 2006). Additionally,
the ventral mPFC is involved with processing intero-
ceptive information important for understanding the
affective significance of stimuli (Decety & Michalska,
2010).
Third, another region implicated in understand-ing others is the inferior frontal gyrus (IFG). The
IFG is part of a set of regions involved with action
understanding and its role in empathy may be in
predicting and understanding action sequences. The
IFG is thought to be part of empathy in regard to
emotional contagion processes, possibly due to simu-
lating others’ emotional states and feeling those same
emotions (Preston & de Waal, 2002; Shamay-Tsoory,
Aharon-Peretz, & Perry, 2009). In addition, the IFG is
important in attentional processes. The IFG, especially
8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 4/12
NEURAL RESPONSES TO SUFFERING 3
right-lateralized, is active in attentional tasks involving
detecting important stimuli (Hampshire, Chamberlain,
Monti, Duncan, & Owen, 2010). The IFG along with
the inferior parietal cortex was more active for empa-
thy when it was elicited via pictures of suffering rather
than cues indicating another was suffering (Lammet al., 2011) and is also active in tasks involving
mentalizing based on perceptual information (e.g.,
Adams et al., 2010), which indicates the importance
of this region in empathy tasks based on perceiv-
ing another’s suffering. In addition, IFG activation is
higher when watching closer friends suffer, mediat-
ing the relationship between friendship and empathy-
related responses in the dACC, which may indicate the
importance of the IFG in greater levels of empathy
for those with whom we are closer (Beeney, Franklin,
Levy, & Adams, 2011).
Despite the fact that almost all of the prior research
on empathy has examined empathy for human suffer-ing, there is some work that has examined empathy
for suffering for animals. Plous (1993, 2003) has
examined human responses to suffering in animals,
especially how it is relevant to whether humans use
animals for human gain. In this research, Plous (1993)
found that the degree to which people perceive similar-
ity with an animal species affects how much empathy
is attributed to suffering animals. Animals consid-
ered more similar to humans were judged as being
more capable of perceiving pain. Further, when par-
ticipants viewed suffering animals, those animals who
were judged as more similar to humans elicited greater
skin conductance responses, indicating that perceiving
suffering in those animals aroused more anxiety in par-
ticipants than perceiving suffering in animals judged to
be less similar to humans.
To our knowledge, only one prior study has com-
pared neural differences in the perception of suffering
in animals and humans. Filippi et al. (2010) exam-
ined responses of vegans, vegetarians, and omnivores
perceiving negatively valenced images of injured and
dead animals in food processing contexts comparing
these to threatening images of violence in humans. For
human suffering, more activation was apparent in the
middle temporal gyrus and precuneus, while animalsuffering led to more activation in the IFG. The ACC
was involved in perceiving suffering in both humans
and animals. Vegetarians and vegans also had greater
responses to animal suffering in several areas involved
with empathy, including the ACC, mPFC, and amyg-
dala. This study examined empathy for animals in
the context of explicit pictures of physical suffering
involving food processing and dietary preferences and
thus could reflect highly charged beliefs and regula-
tory processes due to complicity in the suffering of
animals involved in food processing. In the present
study, we attempted to control for these by examining
whether different neural responses are engaged when
viewing the suffering of humans and dogs, which are
animals we typically engage in a social rather than
dietary manner.
The present research
In our study, we examined neural differences while
people viewed images of humans and dogs suffering.
We chose to use dogs because people are regularly
exposed to dogs and tend to assume that dogs have
greater cognitive abilities than many other animals
(Serpell, 1986). In many ways, humans anthropomor-
phize dogs, extending processes used in understanding
other people to how they understand dogs. People
show high consensus when rating how well Big Five
personality traits apply to pictures of dogs, suggesting
shared implicit knowledge as to how they are assess-
ing dogs’ personalities (Gosling, Kwan, & John, 2003;
Kwan, Gosling, & John, 2008). Further, people also
extend appearance stereotypes found in humans to
their perceptions of dogs, rating more attractive dogs
as having more positive traits and rating more baby-
ish appearing dogs as being more childlike (Zebrowitz
et al., 2011). Dogs may also have a unique abil-
ity among animals to understand humans as well,
as there is evidence that the domestication of dogs
led them to better read human affective states (Hare,Brown, Williamson, & Tomasello, 2002). Darwin even
ascribed human affective states to his dog—such as
joy, guilt, and disappointment—and did not doubt the
“communication and empathy” present between him
and his dog (see Buck & Ginsburg, 1997). Based on
human familiarity and beliefs about the cognitive abili-
ties of dogs, we believed that perceiving dogs suffering
would likely evoke the most powerful responses of
empathy in humans compared to other animals, partic-
ularly those that are commonly associated with dietary
consumption in this culture.
In this study, we examined contrasting predictions
regarding whether the perception of suffering animalsand humans would elicit similar or distinct neural
responses related to empathy. On the one hand, anec-
dotal evidence and anthropomorphism suggests that
we show high levels of empathy for animals, which
leads to the prediction that little difference would be
found in the empathic responses elicited in people who
are viewing animals and human suffering. On the other
hand, empathy research suggests that the degree to
which people empathize with others varies as a func-
tion of perceived similarity they have to the self, which
8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 5/12
4 FRANKLIN ET AL.
might lead one to expect that empathizing with other
humans would activate different neural networks than
when viewing suffering animals.
In addition, one possible reason for differences
in empathy in perceiving suffering in animals and
humans is the possibility that animals elicit greaterempathy due to their perceived helplessness. Empathy-
related responses, such as providing help, are related
to the perceived helplessness of a suffering victim
(Batson et al., 1997; Weiner, 1980). It is possible that
suffering dogs may be seen as more helpless than suf-
fering humans, who by their very nature may be more
responsible for the situations that place them in suffer-
ing. We examined empathy-related neural activation to
suffering children and adults along with suffering adult
dogs and puppies in order to examine this question,
under the supposition that suffering children would be
seen as helpless victims and if helplessness is respon-
sible for differences in neural activation in perceivingsuffering in humans versus animals, these differences
would be weaker in comparisons between children and
puppies than in comparisons between adult dogs and
humans.
To investigate these questions, we examined neu-
ral responses of participants as they perceived images
of suffering humans and dogs. We predicted that per-
ceiving suffering for humans would elicit activation
in brain regions previously implicated in empathy
for the suffering of others, particularly the ACC and
AI. However, as mentioned above, we had compet-
ing predictions as to whether these responses would
extend to animals. Further, we predicted that viewing
suffering in humans compared to animals would acti-
vate regions involved in understanding others’ men-
tal states, including the mPFC (Mason et al., 2004),
whereas perceiving empathy in animals compared
to humans would more likely involve activation in
regions involved in perceptual simulation, such as the
IFG, as was previously found (Filippi et al., 2010).
METHOD
Participants
Seventeen White participants (9 women, 8 men) partic-
ipated for course credit. Participants were either under-
graduate or graduate students, right-handed, under
25 years of age, had normal or corrected to normal
vision, and were free of any history of neurological
problems. Two other participants were dropped for
excess movement in the scanner (>6 mm over the run
while all other participants had <2 mm of movement
over the run). Participants provided informed consent
before completing the study and all study procedures
were approved by the Pennsylvania State University
Institutional Review Board.
Stimuli
One hundred and twenty images of suffering were
selected, with 80 images of humans and 40 images of
dogs suffering. Forty of the images of humans suffer-
ing were of White individuals while 40 images were of
Black individuals.1 Images were selected from Internet
sources and included a wide variety of potential sit-
uations involving suffering, including an equal repre-
sentation of depictions of starvation, physical pain, or
apparent affective suffering across conditions. Further,
images were selected to be free of explicit displays of
blood or gore to avoid responses due to the images
being highly arousing or disgust responses due to vio-
lent displays. We ensured that each group was matchedon how much the individuals looked toward or away
the camera and how near the bodies of the individ-
uals were to the camera. Further, half of the images
were of mature adult humans and dogs and half were
of immature children and puppies.2
Images were pre-rated by an independent sample
of 17 undergraduates (7 female, 10 male) recruited
from the same participant population. Ratings were
made on how much the target in each picture appeared
to be suffering using a seven point scale, with end-
points defined as 1 = not much suffering to 7 = very
much suffering. Responses ranged from a minimum of
2.72 and maximum of 6.56 ( M = 4.51, SD = 0.76).
There were no significant differences in the ratings
of humans and animals in the images selected for the
fMRI participants t (101) = 1.10, p > .27.
Design and procedure
Participants completed one run of the fMRI task where
they viewed images of suffering humans, and dogs.
Participants were instructed that they would see a
series of blocks of images and to merely pay attention
to the images, with no other instructions given about
1 We included same-race and other-race individuals here in order
to examine if any human-specific empathy responses generalized to
other-race individuals. Research examining infrahumanization indi-
cates that humans may not attribute human-like emotional states to
outgroup individuals (e.g., Leyens et al., 2000), thus suggesting that
empathy foroutgroup sufferingmay be more comparable to empathy
for nonhuman entities.2 In addition to the analysis reported here, we compared neural
responses for perceiving suffering in White individuals versus dogs
and Black individuals versus dogs. In these analyses, we found the
same results as reported here, with activations in all the same areas
reported at the threshold we use for this analysis.
8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 6/12
NEURAL RESPONSES TO SUFFERING 5
the images. Participants passively viewed 35 blocks,
10 blocks each of Black individuals, White individ-
uals, and dogs, and 5 blocks of 16 second baseline
fixation. Human images were blocked by race in order
to explore the potential of any cross-race effects. For
each group, five blocks showed mature humans or dogssuffering and five blocks showed immature humans
or dogs suffering. Each block consisted of four pho-
tos, each shown for 3.5 seconds, with a 0.5 second
inter-trial fixation dividing each of the four stimuli.
Thus, each of the 120 images were shown only once
during the entire paradigm. Photos were shown in
random order within blocks for each participant and
blocks were shown sequentially with no time between
blocks. The order of the blocks was pseudo-randomly
determined in order to ensure that the same condi-
tions were not repeated from block to block, with
nine participants viewing one order, and the remain-
ing eight viewing the reverse order. The total run timewas 9 minutes and 20 seconds.
Scanning parameters and analysis
Functional data were collected using a 3T Siemens
Tim-Trio (Siemens AG, Munich, Germany) using
echo-planar imaging with 1 run of 280 T2∗ images
(TR, 2000 ms; TE, 25 ms; 38 interleaved slices; trans-
verse orientation; 3.5 mm slice thickness; 0.5 mm
gap; and voxel size 3.5 mm3). We used SPM8 soft-
ware to process data (Wellcome Institute, London,
UK) using standard six-parameter realignment, coreg-istration of the mean functional image to each subject’s
T1 anatomical image, segmentation of the anatom-
ical image, normalization of functional images to
the Montreal Neurological Institute (MNI) template
using parameters derived from segmentation, and then
smoothing using an 8 mm FWHM Gaussian kernel.
We analyzed functional data using a mass-
univariate general linear model (GLM) approach. For
each subject, we generated first-level fixed-effects con-
trasts for each of the various conditions minus base-
line activation. These first-level contrast images were
then used as the basis for a second-level random-
effects analysis. Second-level random-effects analysesproduced t -statistic maps which were thresholded at
p < .05 false-discovery rate (FDR) corrected, five
voxel cluster extent.
RESULTS
Overlapping activation
In order to assess whether there were regions that
were mutually active for perceiving both human and
animal suffering, we conducted a conjunction analysis
examining activation that was commonly associated
with both human and animal suffering. In order to
examine the overlap between these two conditions, we
computed a conjunction analysis using each of the con-
trasts to baseline (see Friston, Holmes, Price, Büchel,
& Worsley, 1999). However, in order to ensure thatthe areas significant in the conjunction analysis were
significantly active in each condition separately (i.e.,
human minus baseline fixation as well as dog minus
baseline fixation), we created an inclusive mask for
the conjunction analysis for areas that were significant
for both human and animal suffering. To create this
mask, we generated SPM maps of active voxels at the
threshold of p < .05 FDR corrected, five voxel extent
for human suffering minus baseline activation and ani-
mal suffering minus baseline activation and created a
mask of the union of areas significant for both of these
analyses.
This analysis revealed activation in three criticalareas relevant to empathy. First, we found activation
of the bilateral AI and anterior cingulate in both con-
ditions, as well as in the posterior cingulate and cere-
bellum (see Table 1 and Figure 1). This is consistent
with the notion that perceiving animal suffering elicits
empathy-related responses in a similar way as perceiv-
ing human suffering. Second, we found activation of
the mPFC and the IFG. These are regions of the brain
thought to be involved in mentalizing and shown to be
activated in perception of pain in humans. Importantly,
here, we find these areas to be activated when people
perceive pain in both animals and humans. Although
we found differences in the recruitment of these two
areas for human versus animal suffering in the compar-
isons reported above, their conjunction here suggests
that they reflect predominant, though still overlapping
responses.
Differences between humans and dogs
In order to examine differences in suffering between
humans and animals, we conducted a three (type
of sufferer: Black individual, White individual, ordog) within-subjects analysis of variance (ANOVA).
We used this analysis to examine planned contrasts
evaluating differences in suffering for humans versus
animals.
Overall, this ANOVA revealed significant interac-
tions in a network of regions involved with empathy,
including the bilateral parietal cortex, precuneus, and
lateral prefrontal cortex, as well as the left AI, the
bilateral premotor cortex, and dorsal and ventral pre-
frontal cortex (see Table 2). In order to examine
differences in empathy due to race or perceived age of
8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 7/12
6 FRANKLIN ET AL.
TABLE 1
Regions of common activation for human and animal
suffering. Peaks and t -values reflect significant peaks for the
conjunction between the three analyses to baseline
Region x y z t-Value
L. striate cortex −12 −102 8 20.69
R. striate cortex 22 −86 −14 12.41
L. cerebellum −10 −76 −46 5.6
R. cerebellum 26 −76 −42 5.11
R. extrastriate cortex 44 −74 −14 19.33
L. extrastriate cortex −40 −74 −12 15.65
R. precuneus 28 −66 38 4.79
R. occipito-temporal sulcus −34 −64 24 5.2
R. occipito-temporal sulcus 46 −60 18 6.72
L. precuneus −20 −60 44 4.13
Posterior cingulate 0 −54 12 5.35
R. fusiform gyrus 38 −50 −18 11.85
L. fusiform gyrus −40 −44 −24 10.12
L. thalamus −22 −30 −4 7.32
R. parahippocampal gyrus 28 −
26 −
6 7.08R. insula 32 −18 14 2.74
R. precentral gyrus 52 −16 56 4.36
L. precentral gyrus −56 −14 48 4.23
R. superior temporal sulcus 48 −10 −18 2.59
L. amygdala −30 −6 −20 7.96
R. amygdala 28 −6 −16 7.41
L. putamen −24 4 −6 4.2
L. inferior frontal gyrus −36 6 24 4.6
R. inferior frontal cortex 38 8 26 4.56
R. anterior cingulate −4 16 60 3.82
R. temporal pole 44 18 −36 5.07
L. anterior insula −26 18 −22 4.94
L. temporal pole −46 20 −34 6.73
L. ventrolateral prefrontal cortex −54 30 16 4.36
R. orbitofrontal cortex 32 32 −20 4.53
R. vent rolateral prefront al cort ex 58 32 12 3.14
R. ventromedial prefrontal cortex 6 48 −20 5.02
L. ventromedial prefrontal cortex −14 62 −8 3.15
L. dorsomedial prefrontal cortex −10 64 24 5.1
R. dorsomedial pre frontal cortex 10 64 26 4.92
individuals, we conducted exploratory post-hoc com-
parisons examining differences in perceiving suffering
in Black versus White individuals, finding no signifi-
cant differences. In addition, we examined differences
between perceiving human and animal suffering.
First, we compared differences between responses
to humans versus animals (see Table 3 and Figure 2),
using a planned contrast examining differencesbetween humans and animals. We did this using
weighted second-level contrasts in order to account
for having twice as many human stimuli as animal
stimuli.3 Responses to human minus dog suffering
revealed activation in dorsal and ventral regions of
the medial PFC and cerebellum. This likely reflects
the activation of mentalizing processes involved with
3 See note 1.
3 t -value 8
Figure 1. Regions significantly active for suffering in Black and
White individuals as well as for animal suffering compared to
baseline. t -Maps reported reflect the conjunction between contrasts
comparing suffering in each of these groups to baseline with an
inclusive mask for only regions significantly active for all three
conditions to baseline. t -Statistic maps are overlaid on the mean
anatomical image of the participants.
decoding human mental states. Also active wereregions of the posterior cingulate and inferior parietal
lobe, which are involved in self-relevant processing
and thinking in a third-person perspective. Responses
to animal versus human suffering revealed activation
in areas related to affective simulation including the
AI, the premotor cortex, and the IFG. This suggests
that perceiving dog suffering, compared to humans
may capture attention to a greater degree than human
suffering as well as engage more emotional versus
cognitive empathy. Also active were bilateral areas of
the extrastriate cortex and precuneus. These were not
a priori regions of interest, but merit note as they
support evidence asserting that thinking about ani-mals requires more semantic knowledge (Caramazza
& Shelton, 1998) and implies that understanding ani-
mal suffering may involve our semantic knowledge of
animals.
In addition, we examined whether there were any
differences between suffering in immature versus
mature humans and dogs by examining the above-
mentioned ANOVA but three (type of sufferer: Black
individual, White individual, or dog) X 2 (age of suf-
ferer: immature versus mature). This analysis revealed
8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 8/12
NEURAL RESPONSES TO SUFFERING 7
TABLE 2
Regions significant for the ANOVA comparing suffering for White and Black humans, and dogs. Beta
values reflect the beta value for the each contrast at the indicated peak voxel
MNI coordinates Peak beta value
Region x y z F-Value White Black Dog
L. cuneus −8 −92 16 9.92 0.73 1.11 0.81
R. cuneus 16 −90 0 10.63 2.48 3.03 3.25
L. striate cortex −36 −82 18 28.04 0.89 0.97 1.55
R. extrastriate cortex 36 −78 6 30.05 1.53 1.61 2.13
L. extrastriate cortex −34 −76 8 28.66 0.41 0.47 0.81
L. precuneus −18 −70 52 29.79 0.37 0.49 1.42
R. inferior parietal lobule 44 −70 42 11.14 −0.31 −0.24 −0.57
L. inferior temporal cortex −52 −68 −6 33.93 0.75 0.89 1.74
R. precuneus 18 −66 60 28.3 0.06 0.35 1.45
L. angular gyrus −42 −66 48 13.8 −0.26 −0.23 −0.80
R. inferior temporal gyrus 52 −60 −10 34.57 1.26 1.22 2.14
Precuneus 0 −58 30 11.57 0.75 0.90 0.29
R. fusiform gyrus 28 −34 −24 9.28 0.92 0.80 1.10
R. cerebellum 22 −30 −26 8.41 0.46 0.34 0.60
L. postcentral gyrus −60 −26 42 30.8 −0.07 0.16 0.54
R. inferior temporal gyrus 48 −8 −26 10.38 0.19 0.15 0.02
L. insula −46 0 6 13.75 −0.30 −0.03 −0.02
L. premotor cortex −44 2 24 15.8 0.35 0.29 0.68
R. premotor Cortex 48 10 26 16.59 0.73 0.55 1.10
R. dorsomedial PFC 16 40 22 9.35 −0.04 −0.02 −0.14
R. ventromedial PFC 4 48 −18 9.94 1.23 1.20 0.77
R. medial PFC 2 62 4 11.88 1.44 1.45 0.78
no activation at our threshold for the main effects of
age of sufferer the interaction between type of sufferer
and age of sufferer.
DISCUSSION
Anecdotal evidence suggests that we anthropomor-
phize animals and empathize with the suffering with
animals. However, psychological research indicates
that we may think of humans as special. Here, we
found that many of the same brain regions known to be
involved in human empathy are active when perceiving
both human and animal suffering. This suggests that
empathy is not simply a response we keep for humans
alone, but can also extend to nonhuman entities with
which we are familiar. In this study, we found that per-
ceiving suffering in animals, like humans, resulted inthe activation in the primary neural regions known to
be involved in empathy for the perception of suffering
of others, including the ACC and AI.
Empathy for humans versus animals
Despite the similarities in neural responses to human
and animal suffering, when we directly compared
images of suffering humans and animals, we did
find evidence for some distinct patterns of activa-
tion in response to depictions of humans versus ani-
mals, suggesting that different neural mechanisms may
underlie how we derive our empathic responses tohumans and animals. Empathy for humans appears
to elicit more activation in brain regions known to
be involved in decoding others’ mental states, specifi-
cally the mPFC. We found activation in both the dorsal
and ventral mPFC for human versus animal suffer-
ing. Previous work has shown different regions in the
mPFC serve different purposes when decoding others’
mental states. The dorsal mPFC is thought to involve
the use of stereotypical knowledge to understand what
others are thinking while the ventral regions of the
mPFC are involved in understanding similar others
(Mitchell, Macrae, & Banaji, 2006) as well as emo-
tional perspective taking (Hynes et al., 2006). Sucha dissociation indicates the ventral mPFC is involved
in decoding the emotions of others while more dor-
sal regions are involved in understanding meaning
through the actions of others (Amodio & Frith, 2006).
In addition, differences in activation in the ventral
mPFC may reflect differences in concern regarding
perceiving suffering in humans and animals. Lesions to
the ventral mPFC lead to deficits in empathy responses
(Rankin et al., 2006; Sturm et al., 2006). Further,
the development of the mPFC through childhood into
8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 9/12
8 FRANKLIN ET AL.
TABLE 3
Peaks of regions active for the direct contrast between
human and animal and animal versus human empathy
Region x y z t-Value
Human Minus DogL. Inferior Parietal Lobul −44 −74 40 5.91
R. Inferi or Pari etal Lobul e 56 −66 28 5.36
R. Ventromedial Prefront al Cortex 2 62 4 4.85
Posterior Cingulate 0 −58 30 4.58
R. Inferi or Temporal Gyrus 48 −8 −26 4.53
R. Cerebellum 44 −72 −50 4.3
R. Dorsomedial Prefront al Cortex 16 42 22 4.27
L. Inferior Temporal Gyrus −58 −30 −16 4.25
Dog Minus Human
R. Inferi or Temporal Gyrus 52 −60 −10 8.25
L. Inferior Temporal Gyrus −52 −68 −6 8.23
R. Precuneus 18 −60 52 8.2
R. Extrastriate Cortex 36 −78 6 7.75
L. Precuneus −18 −70 52 7.72
L. Extrastriate Cortex −
34 −
76 8 7.76L. Postcentral Gyrus −60 −26 42 7.55
L. Parahippocampal Gyrus −28 −52 −10 7.11
R. Parahippoca mpal Gyrus 28 −50 −14 6.69
R. Postcentral Gyrus 32 −48 68 6.3
R. Inferior Frontal Gyrus 48 8 26 5.37
R. Premotor Cortex 28 −2 52 3.87
R. Cerebellum 40 −44 −42 3.68
L. Striate Cortex −10 −76 10 3.07
R. Lateral Prefrontal Cortex 50 44 10 3.07
R. Thalamus 18 −22 4 2.96
R. Insula 38 −6 −8 2.94
L. Insula −38 −10 −4 2.76
L. Thalamus −14 −24 −2 2.58
L. Inferior Frontal Cortex −44 2 24 5.5
adulthood indicates the importance of this region
in evaluating and regulating responses involved with
empathy (Decety & Michalska, 2010). This hypoth-
esis is also supported as empathy for humans also
evoked more activation in inferior parietal regions and
the posterior cingulate. These regions are implicated in
taking a third-person perspective of others’ situations
(Ruby & Decety, 2001) and distinguishing between
self-produced emotions and emotions as produced by
others (Decety & Grèzes, 2006). Thus, activation in the
ventral mPFC, inferior parietal lobe, and posterior cin-
gulate may represent taking a third-person perspectiveof those who are suffering and distinguishing between
one’s own emotions and the emotions of those who are
suffering.
Observing dog versus human suffering, on the
other hand, led to more activation in a different set
of brain regions, including the AI and IFG. The AI
is important in the affective nature of empathy, or
the feelings elicited by empathy-producing situations,
which suggests that perceiving animal suffering elic-
its greater emotional responses than human suffering,
Human–dog
Dog–human
3 8t -value
Figure 2. Regions active to human minus dog and dog minus
human suffering.
which may form the basis for empathy for animal suf-
fering. Also active for dog versus human suffering
were the IFG and precuneus, replicating findings by
Fillipi et al. (2010). The IFG is active in mentalizingand empathy-related tasks using picture-based stim-
uli (Adams et al., 2010; Lamm et al., 2011). This
may reflect the role of the IFG in action understand-
ing, as perceiving dog suffering may require a greater
degree of understanding the actions of those involved
than perceiving human suffering, which can use more
perspective-taking. Another possibility is that dog suf-
fering captures attention to a greater degree than
human suffering. The IFG is involved in attention allo-
cation as part of the ventral attention system and is
important in allocating attention upon detecting salient
stimuli and unexpected changes in the environment
(Corbetta, Kincade, Ollinger, McAvoy, & Shulman,2000). Participants may have allocated more attention
to animal suffering because they likely do not have
as much experience with animal suffering or because
our participants were not given specific commands in
regard to the stimuli. This is especially notable as a
right-lateralized ventral attention system is thought to
be involved even without specific attentional demands
(Fox, Corbetta, Snyder, Vincent, & Raichle, 2006).
Interestingly, we did not find differences in neu-
ral responses when perceiving suffering children and
8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 10/12
NEURAL RESPONSES TO SUFFERING 9
puppies as compared to adult dogs and humans. This
suggests that the differences we found between human
and animal suffering were not due to relative differ-
ences in how helpless humans and dogs were perceived
due to their suffering. However, the lack of differ-
ences between perceiving suffering children and adultsis interesting in the light that children should evoke
greater empathy than adults because they are seen as
more helpless. One limitation of our study is that we
showed a series of still images of suffering, whereas it
is more ecologically valid to see suffering as a single
presentation. It is possible that more ecologically valid
presentations, such as high-definition video, would
lead to greater differences in perceiving suffering in
animals and humans.
CONCLUSIONS
The present study indicates that there are many over-
lapping regions in humans’ empathic responses to
viewing animal and human suffering, particularly in
areas classically associated with empathic response.
Direct comparisons also revealed different neural sub-
strates activated for empathy for humans than empathy
for animals. This finding is important as it suggests
that the way we arrive at our empathic responses
to the suffering of nonhuman entities, in this case
dogs, may be fundamentally different than how we
arrive at similar responses to suffering in humans.
Nonetheless, many of the regions that were differen-
tially active when comparing human versus animalsuffering were nonetheless significantly active for both
conditions in perceptions of suffering to baseline, indi-
cating that although they represent potentially different
sources for generating similar empathic responses,
they are not mutually exclusive, but rather differen-
tially predominant.
One notable limitation of our research is that we
only examined a homogenous college-aged popula-
tion. This population is relatively protected when it
comes to perceiving suffering animals, especially in
the context of worldwide and historical interactions
between humans and animals. It is possible these find-
ings may not extend to other individuals who are more
experienced with perceiving animal suffering or to cul-
tures that use animals more directly as beasts of burden
or for survival.
Future research is necessary to examine the degree
to which these findings generalize to perceptions of
suffering in other animals aside from dogs. The current
research examined empathy-related neural processes
in the perception of suffering in dogs for the purpose
of choosing an animal that would evoke the great-
est degree of human-like empathy processes, due to
human experience with dogs. However, this may indi-
cate that perceiving suffering in other species that
humans see as less like humans or have less expe-
rience with would evoke less empathy-related neuralactivation. As discussed in the introduction, Plous’s
(1993) similarity hypothesis indicates that humans
show greater empathy for judging suffering in ani-
mals perceived as more similar to humans. In addi-
tion, many of the same perceptual processes used for
judging traits in humans extend to how we judge ani-
mals (e.g., Franklin, Zebrowitz, Fellous, & Lee, 2013;
Kwan et al., 2008). For example, Zebrowitz et al.
(2011) found that perceptual stereotypes associated
with babyfaceness extend to perceptions of animals,
with more babyish-looking animals evoking greater
stereotypic associations with being more likeable and
warm. Based on this, we would predict that animalswho we perceive to be more likeable and warm may
elicit greater empathy, but this is a question for future
examination.
Empathy is a critical motivation for prosocial
behavior and as such, understanding the processes that
elicit empathy is essential to understand why and how
we engage in prosocial behavior. Responses to animal
suffering motivate a powerful animal rights movement
(Singer, 2002). This is evident in the rise of vegetari-
anism and veganism practices in dietary habits (Kim,
Schroeder, Houser, & Dwyer, 1999) and increases
in the number of people opposing animal experi-
mentation (National Science Board, 2010). Stories
arousing empathy for animal suffering may increase a
moral focus on the value of nature (Berenguer, 2010).
Additionally, animal suffering may be used to promote
awareness for more broad-based environmental top-
ics such as deforestation and anthropogenic climate
change (Cole et al., 2009). The research presented here
suggests the neural responses underlying empathy for
perceiving animal suffering are highly similar as for
perceiving human suffering, but with some potentially
critical differences in the mechanisms driving how we
arrive at these responses. Many studies detail how
empathy plays a critical role in prosocial behavior tohumans, but it is unclear how empathy for animals may
form a basis for pro-animal behavior. This is a critical
question for future research, especially in the context
of social and political issues that may affect the welfare
of humans and animals differently.
Original manuscript received 23 October 2012
Revised manuscript accepted 2 January 2013
First published online 12 February 2013
8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 11/12
10 FRANKLIN ET AL.
REFERENCES
Adams, R. B. Jr, Rule, N. O., Franklin, R. G., Jr, Wang,E., Stevenson, M. T., Yoshikawa, S., . . . Ambady, N.(2010). Cross-cultural reading the mind in the eyes: AnfMRI investigation. Journal of Cognitive Neuroscience,22, 97–108.
Amodio, D. M., & Frith, C. D. (2006). Meeting of minds:The medial frontal cortex and social cognition. Nature
Reviews Neuroscience, 7 , 268–277.Batson, C. D., Polycarpou, M. P., Harmon-Jones, E., Imhoff,
H. J., Mitchener, E. C., Bednar, L. L., . . . Highberger, L.(1997). Empathy and attitudes: Can feeling for a mem-ber of a stigmatized group improve feelings toward thegroup? Journal of Personality and Social Psychology, 72,105–118.
Beeney, J. E., Franklin, R. G. Jr, Levy, K. N., & Adams,R. B. (2011). I feel your pain: Emotional closenessmodulates neural responses to empathically experiencedrejection. Social Neuroscience, 6 , 369–376.
Berenguer, J. (2010). The effect of empathy in environ-
mental moral reasoning. Environment and Behavior , 42,110–134.
Buck, R., & Ginsburg, B. (1997). Communicative genesand the evolution of empathy. Annals of the New York
Academy of Sciences, 807 (1), 481–483.Caramazza, A., & Shelton, J. R. (1998). Domain-specific
knowledge systems in the brain: The animate–inanimatedistinction. Journal of Cognitive Neuroscience, 10(1),1–34.
Cheng, Y., Chen, C., Lin, C. P., Chou, K. H., & Decety, J.(2010). Love hurts: An fMRI study. NeuroImage, 51(2),923–929.
Cole, M., Miele, M., Hines, P., Zokaei, K., Evans, B.,& Beale, J. (2009). Animal foods and climate change:Shadowing eating practices. International Journal of
Consumer Studies, 33(2), 162–167.Corbetta, M., Kincade, J. M., Ollinger, J. M., McAvoy, M.
P., & Shulman, G. L. (2000). Voluntary orienting is dis-sociated from target detection in human posterior parietalcortex. Nature Neuroscience, 3, 292–297.
Craig, A. D. (2002). How do you feel? Interception: Thesense of the physiological condition of the body. Nature
Reviews Neuroscience, 3, 655–666.Davidson, R. J., Putnam, K. M., & Larson, C. L.
(2000). Dysfunction in the neural circuitry of emotionregulation—a possible prelude to violence. Science, 289,591.
Decety, J. (2007). A social cognitive neuroscience model of human empathy. In E. Harmon-Jones & P. Winkielman(Eds.), Social neuroscience: Integrating biological and
psychological explanations of social behavior (pp.246–270). New York, NY: Guilford Press.Decety, J., & Grèzes, J. (2006). The power of simula-
tion: Imagining one’s own and other’s behavior. Brain Research, 1079(1), 4–14.
Decety, J., & Michalska, K. J. (2010). Neurodevelopmentalchanges in the circuits underlying empathy and sympathyfrom childhood to adulthood. Developmental Science, 13,886–899.
Eisenberger, N. I., & Lieberman, M. D. (2004). Why rejec-tion hurts: A common neural alarm system for physi-cal and social pain. Trends in Cognitive Sciences, 8(7),294–300.
Eslinger, P. J. (1998). Neurological and neuropsychologicalbases of empathy. European Neurology, 39(4), 193–199.
Filippi, M., Riccitelli, G., Falini, A., Di Salle, F.,Vuilleumier, P., Comi, G., & Rocca, M. A. (2010). Thebrain functional networks associated to human and ani-mal suffering differ among omnivores, vegetarians and
vegans. PLoS One, 5(5), e10847.Fox, M. D., Corbetta, M., Snyder, A. Z., Vincent, J. L., &
Raichle, M. E. (2006). Spontaneous neuronal activity dis-tinguishes human dorsal and ventral attention systems.Proceedings of the National Academy of Sciences, 103,10046–10051.
Franklin, R. G. Jr, Zebrowitz, L. A., Fellous, J. -M., & Lee,A. (2013). Human ability and cultural variation in sex-differentiating macaque faces. Journal of ExperimentalSocial Psychology, 49, 344–348.
Friston, K. J., Holmes, A. P., Price, C. J., Büchel, C., &Worsley, K. J. (1999). Multisubject fMRI studies andconjunction analyses. NeuroImage, 10, 385–396.
Gosling, S. D., Kwan, V. S., & John, O. P. (2003). A dog’sgot personality: A cross-species comparative approach to
personality judgments in dogs and humans. Journal of Personality and Social Psychology, 85, 1161–1169.
Hampshire, A., Chamberlain, S. R., Monti, M. M., Duncan,J., & Owen, A. M. (2010). The role of the right infe-rior frontal gyrus: Inhibition and attentional control.
NeuroImage, 50, 1313–1319.Hare, B., Brown, M., Williamson, C., & Tomasello, M.
(2002). The domestication of social cognition in dogs.Science, 298(5598), 1634.
Holroyd, C. B., Nieuwenhuis, S., Yeung, N., Nystrom, L.,Mars, R. B., Coles, M. G. H., . . . Cohen, J. D. (2004).Dorsal anterior cingulate cortex shows fMRI response tointernal and external error signals. Nature Neuroscience,7 , 497–498.
Hynes, C. A., Baird, A. A., & Grafton, S. T. (2006).Differential role of the orbital frontal lobe in emotionalversus cognitive perspective-taking. Neuropsychologia,44(3), 374–383.
Kennedy, J. S. (1992). The new anthropomorphism. NewYork, NY: Cambridge University Press.
Kim, E. H. J., Schroeder, K. M., Houser, R.F., & Dwyer, J. T.(1999). Two small surveys, 25 years apart, investigatingmotivations of dietary choice in 2 groups of vegetari-ans in the Boston area. Journal of the American Dietetic
Association, 99(5), 598–601.Kohn, A. (1990). The brighter side of human nature:
Altruism and empathy in everyday life. New York, NY:Basic Books.
Kwan, V. S. Y., Gosling, S. D., & John, O. P. (2008).Anthropomorphism as a special case of social perception:A cross-species social relations model analysis of humans
and dogs. Social Cognition, 26 , 129–142.Lamm, C., Decety, J., & Singer, T. (2011). Meta-analytic
evidence for common and distinct neural networks asso-ciated with directly experienced pain and empathy forpain. NeuroImage, 54, 2492–2502.
Leyens, J. P., Paladino, P. M., Rodriguez-Torres, R., Vaes, J.,Demoulin, S., Rodriguez-Perez, A., & Gaunt, R. (2000).The emotional side of prejudice: The attribution of sec-ondary emotions to ingroups and outgroups. Personalityand Social Psychology Review, 4(2), 186.
Mason, M. F., Banfield, J. F., & Macrae, C. N. (2004).Thinking about actions: The neural substrates of personknowledge. Cerebral Cortex , 14(2), 209–214.
8/13/2019 Perceived Pain in Animals
http://slidepdf.com/reader/full/perceived-pain-in-animals 12/12
NEURAL RESPONSES TO SUFFERING 11
Mitchell, J. P., Macrae C. N., & Banaji, M. R. (2006).Dissociable medial prefrontal contributions to judgmentsof similar and dissimilar others. Neuron, 50(4), 655–663.
National Science Board. (2010). Science and engineer-ing indicators: 2010. Arlington, VA: National ScienceFoundation.
Ostrowsky, K., Magnin, M., Ryvlin, P., Isnard, J., Guenot,M., & Mauguière, F. (2002). Representation of painand somatic sensation in the human insula: A studyof responses to direct electrical cortical stimulation.Cerebral Cortex , 12(4), 376.
Plous, S. (1993). Psychological mechanisms in the humanuse of animals. Journal of Social Issues, 49, 11–52.
Plous, S. (2003). Is there such a thing as prejudice towardanimals? In S. Plous (Ed.), Understanding prejudiceand discrimination (pp. 509–528). New York, NY:McGraw-Hill.
Preston, S. D., & de Waal, F. B. M. (2002). Empathy:Its ultimate and proximate bases. Behavioral and BrainSciences, 25, 1–20.
Rankin, K. P., Gorno-Tempini, M. L., Allison, S. C., Stanley,
C. M., Glenn, S., Weiner, M. W., & Miller, B. L. (2006).Structural anatomy of empathy in neurodegenerative dis-ease. Brain, 129, 2945–2946.
Ruby, P., & Decety, J. (2001). Effect of subjective per-spective taking during simulation of action: A PETinvestigation of agency. Nature Neuroscience, 4(5), 546.
Serpell, J. (1986). In the company of animals. Oxford: BasilBlackwell.
Shamay-Tsoory, S. G., Aharon-Peretz, J., & Perry, D.(2009). Two systems for empathy: A double dissocia-tion between emotional and cognitive empathy in inferiorfrontal gyrus versus ventromedial prefrontal lesions.
Brain, 132, 617–627.Singer, P. (1974). All animals are equal. Philosophical
Exchange, 1(Summer), 103–116.Singer, P. (1975). Animal liberation: A new ethics for our
treatment of animals. New York, NY: Random House.Singer, T., Critchley, H. D., & Preuschoff, K. (2009).
A common role of insula in feelings, empathy,and uncertainty. Trends in Cognitive Sciences, 13(8),334–340.
Singer, T., Seymour, B., O’Doherty, J., Kaube, H., Dolan,R. J., & Frith, C. D. (2004). Empathy for pain involvesthe affective but not sensory components of pain. Science,303, 1157.
Sturm, V. E., Rosen, H. J., Allison, S., Miller, B. L.,& Levenson, R. W. (2006). Self-conscious emotiondeficits in frontotemporal lobar degeneration. Brain, 129,2508–2516.
Weiner, B. (1980). A cognitive (attribution)-emotion-actionmodel of motivated behavior: An analysis of judg-ments of help-giving. Journal of Personality and SocialPsychology, 39, 186–200.
Zebrowitz, L. A., Wadlinger, H. A., Luevano, V. X., White,B. M., Xing, C., & Zhang, Y. (2011). Animal analo-gies in first impressions of faces. Social Cognition, 29,486–496.