Why Don’t Men Understand Women? Altered Neural
Networks for Reading the Language of Male and Female
Eyes
Boris Schiffer1,2*, Christina Pawliczek2,5, Bernhard W. Müller3, Elke R. Gizewski4,6, Henrik Walter7
1 Division of Forensic Psychiatry, Department of Psychiatry, Psychotherapy and Preventive Medicine, LWL-University Hospital Bochum, Germany, 2 Institute of Forensic
Psychiatry, University of Duisburg-Essen, Germany, 3 Department of Psychiatry and Psychotherapy, University of Duisburg-Essen, Germany, 4 Department of Diagnostic
and Interventional Radiology and Neuroradiology, University Hospital Essen, Germany, 5 Department of Psychiatry, Psychotherapy, and Psychosomatics, University
Hospital Aachen, RWTH Aachen, Germany, 6 Department of Neuroradiology, University Hospital Innsbruck, Austria, 7 Division of Mind and Brain Research, Department of
Psychiatry and Psychotherapy, Charité Universitätsmedizin Berlin, Berlin, Germany

Abstract
Men are traditionally thought to have more problems in understanding women compared to understanding other men,though evidence supporting this assumption remains sparse. Recently, it has been shown, however, that mens problems inrecognizing women’s emotions could be linked to difficulties in extracting the relevant information from the eye region,
which remain one of the richest sources of social information for the attribution of mental states to others. To determine
possible differences in the neural correlates underlying emotion recognition from female, as compared to male eyes, amodified version of the Reading the Mind in the Eyes Test in combination with functional magnetic resonance imaging(fMRI) was applied to a sample of 22 participants. We found that men actually had twice as many problems in recognizing
emotions from female as compared to male eyes, and that these problems were particularly associated with a lack ofactivation in limbic regions of the brain (including the hippocampus and the rostral anterior cingulate cortex). Moreover,men revealed heightened activation of the right amygdala to male stimuli regardless of condition (sex vs. emotion
recognition). Thus, our findings highlight the function of the amygdala in the affective component of theory of mind (ToM)
and in empathy, and provide further evidence that men are substantially less able to infer mental states expressed by
women, which may be accompanied by sex-specific differences in amygdala activity.
Citation: Schiffer B, Pawliczek C, Müller BW, Gizewski ER, Walter H (2013) Why Don’t Men Understand Women? Altered Neural Networks for Reading theLanguage of Male and Female Eyes. PLoS ONE 8(4): e60278. doi:10.1371/journal.pone.0060278
Editor: Frank Krueger, George Mason University/Krasnow Institute for Advanced Study, United States of America
Received November 14, 2012; Accepted February 24, 2013; Published April 10, 2013
Copyright: ß 2013 Schiffer 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: This research was funded by the Landschaftsverband Rheinland, Germany (Dr. Schiffer). The funders had no role in study design, data collection andanalysis, decision to publish, or preparation of the manuscript.
Competing Interests: The authors have declared that no competing interests exist.
* E-mail: boris.schiffer@uni-due.de

Recent studies have suggested that deficits in recognizing facial
emotions could be linked to difficulties in extracting the relevant
information from the eye region (including gaze direction) [12],which remains one of the richest sources of social information for
the attribution of mental states to others. Recent evidence has
further indicated that social cognitive processes such as faceprocessing and mental state attribution continue to develop duringadolescence [13], and are due to genetic and/or hormonal (e.g.
fetal testosterone) effects on brain maturation, which mightdevelop distinctively in boys and girls [14,15].
The Reading-the-Mind-in-the-Eyes Test (RMET) is based on
photographs of isolated eye regions, and has been developed forthe evaluation of mentalizing capacities in adults [16] andchildren [17]. In this test, four words indicating an emotional
mental state are presented and subjects are required to choose
which one best describes the emotion expressed in the eyeregion. A recent study using this test in a sample of university
students has shown that men performed better in recognizing
emotions from male vs. female eyes under placebo vs.
vasopressin administration [2].

Introduction
Sex differences in brain morphology and cognitive and affectivefunctions have been increasingly documented [1]. One study has
found that men have more problems in recognizing mental statesand emotions in women than in men [2], and it is still unclear why
this could be. Other studies have found that relative to women
men are faster in detecting angry male faces [3]. Some authors
have suggested that same-sex facial expressions may be more
important to males as compared to females [4].
A so-called theory of mind (ToM) network and an empathy
network of brain regions have been found to underlie the processes
required to understand and share other people’s thoughts, feelings
and intentions. The ToM network comprises of the following
regions: the precuneus, posterior superior temporal sulcus (pSTS),
temporo-parietal junction (TPJ), temporal poles, and medial
prefrontal cortex (mPFC) [5,6], whereas the empathy network
includes the following brain areas: the anterior cingulate cortex(ACC) and the anterior insula [7]. However, the amygdala, whichis also implicated in face processing [8,9], also seems to play animportant role in emotional empathy [10] and affective ToM [11].

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Based on these results, we expected that better performance inthe recognition of emotions in male vs. female eyes would be
related to differential (stronger) recruitment of networks underlying emotional empathy or affective ToM (i.e. the amygdala,anterior insula, and anterior cingulate cortex) [7,10]. We
examined a sample of healthy male participants using fMRI in
combination with a modified version of the RMET [17], which
required both sex discrimination and emotion recognition of thesame pairs of female and male eyes.

All MR images were obtained using a 1.5 T MR (Sonata,
Siemens, Erlangen, Germany) with a standard head coil. BOLD
contrast images were acquired by applying an echo-planaracquisition technique (TR 3500 ms, TE 45 ms, flip angle 90u,
FOV 240 mm, matrix 64) with 38 transversal slices (thickness
3.863.863 mm) and a 0.3 mm slice gap. Six initial ‘‘dummy’’scans were eliminated prior to the data analysis to account for T1
relaxation effects.

Materials and Methods

Image Processing

Data Acquisition

We used SPM8 software (http://www.fil.ion.ucl.ac.uk/spm/)
for the analysis of the imaging data. Prior to second level statisticalanalyses, the images were realigned using sinc interpolation andnormalized to the stereotactic template of the Montreal Neurological Institute (http://imaging.mrc-cbu.cam.ac.uk/imaging/
MniTalairach). Bilinear interpolation was applied for normalization to the MNI-template. Normalized images were smoothedwith an isotropic Gaussian kernel of 9 mm FWHM. Single subject
contrasts between experimental and control conditions werecomputed. The model consisted of a boxcar function convolved
with the hemodynamic response function (HRF) and thecorresponding temporal derivative [26]. High-pass filtering witha cutoff frequency of 120 sec. and low-pass filtering with the HRF
were applied.

Participants
22 single men aged between 21 and 52 years (mean age:
35.6610.0) old were recruited for participation through advertisements. All participants were right-handed, medically andpsychiatrically healthy, and with an estimated IQ [18] greaterthan 80 (mean IQ: 109.8612.1). None of the subjects had any
history of psychiatric illness, significant physical medical conditions, or any other condition that would interfere with MRI
scanning (e.g., extreme obesity, claustrophobia, cochlear implant,
metal fragments in eyes, cardiac pacemaker, neural stimulator,and metallic body inclusions or other metal implanted in thebody). Following initial screening, subjects were interviewed about
their medical history. Furthermore, the SCID I [19] and II [20]were administered by an experienced psychiatrist trained to use
these instruments.
The study was approved by the local Committee on Medical
Ethics of the Medical Faculty of the University of Duisburg-Essen,
Germany and was performed in accordance with the Code of
Ethics of the World Medical Association (Declaration of Helsinki).
After a detailed description of the study was given to participants,
written informed consent was obtained.

Statistical Analysis
Behavioral data. We performed a repeated measures analysis of variance (ANOVA) with two within subjects factors: (1)stimulus type (male eyes vs. female eyes) and (2) condition (emotion
vs. gender recognition) in order to confirm that men had greater
problems recognizing emotion than gender ( = main effect ofcondition) as well as to explore whether men reveal greater
problems in recognizing emotions from female as compared tomale eyes ( = condition x stimulus type interaction).
fMRI data. We used the framework of the General Linear
Model to perform a statistical group analysis on the fMRI data on
a voxel-by-voxel basis. Analyses included five mutually exclusive
trial covariates: correct emotion recognition from male and female
eyes as well as correct gender recognition from male and female
eyes and errors (either from emotion or gender recognition trials).
Partial correlation maps for individual participants were generated, indicating the extent to which each voxel’s activity conformed
to an a priori canonical double gamma hemodynamic response
function. Between-condition contrasts (i.e., gender vs. emotion
recognition for all stimuli as well as male and female eyes
separately) were estimated on a voxel-by-voxel basis in a wholebrain approach. T Maps of activations for emotion and genderrecognition in male and female eyes were submitted to randomeffects ANOVAs to test for the main effect of condition (activation
on emotion recognition minus activation on gender recognition)
the main effect of stimulus type (male vs. female eyes), and thecondition by stimulus type interaction among all subjects. Similar
to earlier studies [22,24], statistical thresholds were set to p,.001,
uncorrected at the voxel level (height threshold) and p,.05,
corrected after Family Wise Error (FWE) at the cluster level (extent
threshold) in all whole-brain analyses. To increase sensitivity forstimulus and condition x stimulus interaction effects, we performed
small volume corrections (SVC), if differential activations in theseregions were predicted by our a priori hypothesis. For this purpose
ROIs within areas related to affective empathy (i.e. the amygdala,anterior insula, and anterior cingulate cortex) were a priori definedby using the Automatic Anatomic Labeling (AAL) [27]. Sphericalvolumes with a 10-mm radius were centered on the coordinates offMRI Task
A modified version of the ‘‘Reading the Mind in the Eyes Test’’(RMET)[16,21–24], was administered to all participants in thescanner. The RMET assesses the ability to infer other people’s
mental states, using information from 36 pairs of eyes (black-andwhite pictures of eighteen male and eighteen female eyes). All
stimuli were of equal size (2268 cm) and depicted twelve negative,
eight positive and sixteen neutral expressions, counterbalanced forthe sex of the eyes [25].
Participants were asked to decide which of the two presented
words (e.g., distrustful or terrified) best described the emotional/
mental state of the person whose eyes were presented. The control
condition comprised of a sex discrimination task in which for thesame 36 pictures of eyes, the participants had to judge whether theeyes belonged to a man or woman. In total, the tasks consisted of
72 stimuli presented in a randomized (regarding stimuli withinblocks and block order) block design with 12 blocks of six pairs ofeyes. Participants responded via button press with their right index
and middle fingers.
At the start of each block, the word ‘‘Emotion’’ or ‘‘Gender’’was presented for seven seconds. The blocks assessing mental
states were alternated with those assessing gender. Each picture
was presented for eight seconds, and a question mark presented in
the sixth second prompted the subjects to answer. Each block
therefore lasted 48 seconds. Behavioral data were acquired with abutton pad (LumitouchTM Photon Control Inc, Burnaby, Canada). RMET scores were calculated as the percentage of correct
discriminations over conditions, eye- type, and emotional valence
as well as response times after the six-seconds period.

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triangularis; extending into the anterior insula and temporal pole;
MNI: 250, 28,28; pFWEc,.001; z = 6.03; k = 3182), the bilateral
pSTS/TPJ (MNI: 258, 242,6; pFWEc,.001; z = 5.78; k = 1407;
MNI: 52, 236,22; pFWEc = 0.032; z = 4.70; k = 200), and themPFC/ACC (MNI: 22, 20,54; pFWEc = 0.048; z = 4.36; k = 160).
There were also activations in the bilateral occipital cortex (MNI:
216, 298,28; pFWEc = 0.037; z = 4.06; k = 192; MNI: 32,
290,212; pFWEc = 0.173; z = 3.87; k = 118), of which the rightcluster passed the significance threshold at the voxel but not at thecluster level. As illustrated in Fig. 2a, there were significant
condition-by-stimulus-type interactions for activation of the rightprecuneus, right dorsolateral PFC (dlPFC), right hippocampus,
and left rostral ACC (rACC; details regarding coordinates andstatistics are provided in the figure legend). Contrast estimates asprovided in Fig. 2b showed that interaction effects of the
hippocampus and rACC reflect increased activation while
recognizing emotions from male as compared to female eyes,
whereas interaction effects of the precuneus and the dlPFC reflect
the opposite pattern. Interestingly, there was also a significant
main effect for stimulus type in the right amygdala (MNI:
18,24,220, pFWE-SVC = 0.004; z = 4.10; cluster-size = 69), reflecting increased activation to male relative to female eyes regardlessof condition (see Fig. 3).
Since right amygdala activation to male eyes was higher
compared to female eyes regardless of condition, but also seemed
to be descriptively higher during the emotion recognition
condition, we tested whether right amygdala activation modulated
recognition accuracy. We therefore extracted non-adjusted activation (main effect: male vs. female stimuli) of the cluster within
the right amygdala using volume of interest analysis (1steigenvariate) and calculated Pearsons correlation coefficient
between right amygdala responses and recognition accuracy formale and female stimuli separately. There was a significant
positive association between right amygdala activation andthe peak voxel within the a priori defined ROI. Significancethresholds for both height and extent were set to p,.05 FWE
corrected for multiple comparisons as well as the number of ROIs
(362) at the voxel level.
Regression models were calculated to assess the associations ofage with differences in performance accuracy and activation
pattern. Activations in regions showing significant group differences were extracted by volume of interest analyses (1steigenvariate) including all voxels of each cluster. Regression models were
calculated step-wise with F probabilities of ,.05 to enter and ..1
to remove variables. Multi co-linearity was tested by calculating
the squared multiple correlation (R2) of predictor variables.

Results
As illustrated in Fig. 1a, men exhibited significantly greater
problems recognizing emotion than gender (main effect ofcondition: F1,21 = 39.5; p,.001). Furthermore, they showed
greater problems recognizing emotions from female compared tomale eyes (condition x stimulus-type interaction: F1,21 = 24.1;
p,.001). The same result was found for the reaction timemeasures (Fig. 1b). Men exhibited significantly longer reaction
times when recognizing emotions than gender (main effect ofcondition: F1,21 = 16.0; p,.001) and the RT differences (emotion –gender recognition) were significantly larger for female than maleeyes (condition x stimulus-type interaction: F1,21 = 4.5; p = .042).
To confirm that differences in the ability to recognize emotions
from male and female eyes were not affected by emotional valenceof stimuli, an additional two (male vs. female eyes) by three(negative vs. positive vs. neutral expressions) ANOVA was
performed. Neither a significant main effect for emotional valence(F1,21 = 0.54; p = .819) nor a significant eyes type by emotional
valence interaction (F1,21 = 2.83; p = .107) could be detected.
Emotion as compared to gender recognition led to significant
activation of the left inferior frontal gyrus (pars orbitalis andFigure 1. (a) Bar chart illustrating performance accuracy (% correct) and (b) line plot illustrating reaction time measures over conditions (emotion vs.
gender recognition) and types of stimuli (male vs. female eyes). Significance bars and asterisks designate the significance of both recognition
accuracy and reaction times for the main effects of condition and the condition by eyes type interactions.
doi:10.1371/journal.pone.0060278.g001

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Figure 2. (a) Statistical parametric maps and (b) parameter estimates illustrating significant condition–by-stimulus-type interaction effects for the
rACC (MNI: 26,30,210; pFWE-SVC = 0.005; z = 3.85; cluster-size(k) = 194), right hippocampus (MNI: 32,28,226; pFWEc = 0.025; z = 4.37; k = 130), precuneus(MNI: 28,270,36; pFWEc = 0.034; z = 4.54; k = 494), and dlPFC (MNI: 52,32,22; pFWEc = 0.046; z = 3.85; k = 291). All coordinates reference the coordinate
system of the Montreal Neurological Institute (MNI). Color bar indicates z-statistic value.
doi:10.1371/journal.pone.0060278.g002

mentalizing performance for male (r = .59; p = .004), but not
female stimuli (r = -.07; p = .760; see Fig. 3c).
Given that previous research has documented a negative
association between age and mentalizing performance [28], we
explored whether stimulus-related differences in mentalizing
performance and neural responses were linked to age. There
was a negative association between age and mentalizing performance (beta = -.466; p = .029) as well as differential right amygdala
activity (beta = -.419; p = .049). Yet, performing additional analyses of covariance including age as a covariate, neither thecondition-by-stimulus-type interaction on recognition accuracy(F1,20 = 5.0; p = .009) nor differential right amygdala activity(MNI: 18,22,220, z = 3.9) were solely a consequence of its
association with age.

interaction (F6,30 = 0.71; p = .500). Thus, it seems unlikely thatthe finding of different responsiveness of the right amygdala tomale vs. female eyes is the result of differences in pupil diameter
size.

Discussion
In this study, we confirmed recent research documenting that
men actually have greater difficulties to accurately assess
emotional expressions via female as compared to male eyes [2].
In addition, we showed for the first time that better performance inrecognizing emotions from male vs. female eyes was linked todifferential activation patterns in areas that have been previously
linked to mentalizing performance [2,3]. While the main effect ofcondition confirmed previous research documenting that emotion
recognition, relative to gender, recruited areas of the mentalizing
network (including pSTS/TPJ, temporal poles, and mPFC/ACC)
as well as areas predicting the affective consequences of a stimulus(i.e. the anterior insula [7]), the condition-by-stimulus-type
interaction revealed significant effects in four regions (i.e.
hippocampus, rostral ACC, dlPFC, and precuneus), indicating
differential activation while recognizing emotions as compared togender in both types of stimuli (male vs. female). Emotion vs.
gender recognition in male relative to female eyes was associatedPost-hoc Analysis
Since previous research has shown a relationship between
amygdala activity and pupil diameter changes in faces withdifferent emotions [29,30], we analyzed whether pupil diametersize differed due to eye type, emotional valence or both. A two (eyetype) by three (emotional valence) ANOVA on pupil diametersizes, however, revealed neither a significant main effect of eye
type (F6,30 = 2.29; p = .118), or emotional valence (F6,30 = 1.84;
p = .184) nor a significant eye type by emotional valence
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Figure 3. (a) Statistical parametric map and (b) parameter estimates illustrating differential activation pattern of the right amygdala (MNI: 18,24,220,
pFWE-SVC = 0.004; z = 4.10; cluster-size = 69) for male and female eyes (regardless of condition: emotion vs. gender recognition). Color bar indicates zstatistic value. (c) Scatter plots depicting the relationships between amygdala response and emotion recognition accuracy for both types of stimuli.
doi:10.1371/journal.pone.0060278.g003

with females in which the recognition of the specific emotion had
occurred.
The finding of heightened right amygdala responses during
recognition of male compared to female stimuli might indicate ahighly automated and stimulus-driven effect that occurred
regardless of different conditions or instructions. Thus, increased
right amygdala responses to male stimuli may indicate a sexspecific association between stimulus type (male vs. female) andautomated emotion processing or affective empathy. The positive
correlation between right amygdala responses and the ability toinfer mental states from male but not female eyes also indicatedthat affective empathy might enhance mentalizing performancefor male stimuli in men. Conversely, this indicated that theprocessing of opposite-sex stimuli might be associated with lower
affective empathy which may also be associated with a reduced
mentalizing ability. In support of this assumption, a previous study
investigating emotion recognition accuracy in two patients withacquired amygdala lesions showed impaired emotion recognition
using an earlier version of the RMET [38]. Results from this study
particularly showed impairment in the male patient with right
amygdala damage, whereas the female patient with bilateral
amygdala lesion did not present any mentalizing impairments.
Notably, both patients made comparable numbers of errors on
items on the RMET [28].
Moreover, it is possible that the pattern of seemingly decreased
right amygdala (from the main effect of male vs. female eyes) andincreased right dlPFC activation while inferring emotions fromfemale relative to male eyes, reflected a down-regulation
mechanism of emotional experiences specific to female stimuli,
which bears an additional source of information for the attributionof mental states to women. If true, one would expect a negative
correlation between activation of the right dlPFC and theamygdala, which is exactly what we found. We observed asignificant negative correlation of -.438 (p = .042) between
activation of the right dlPFC and right amygdala during emotion
recognition. Thus, this mechanism might account for the relatively
greater problems men have in recognizing women’s compared tomen’s emotions.
Interestingly, intranasal administration of nonapeptide argininevasopressin (AVP) has been shown to negatively affect mens
emotion recognition accuracy to same- but not opposite-sex

with heightened activation in the right hippocampus and therostral ACC, whereas emotion vs. gender recognition in female
relative to male eyes was associated with heightened activation of
the right dlPFC and the right precuneus. Moreover, we found thatright amygdala responses were increased for male stimuli
regardless of condition (emotion vs. gender recognition).
Both the hippocampus and the rACC are involved in theacquisition and expression of emotional memories [31]. Thus,
increased activation of these regions during recognition ofemotions vs. gender in male relative to female eyes may reflect
relatively stronger use of autobiographical memory information
when taking others mens perspective. Furthermore, Perry et al.
reported increased hippocampus activation during an emotional
mentalizing task and suggested that in order to better understandand empathize with others, memories are projected to people thatare seen as similar to oneself [32]. Though they used a different
task, a similar phenomenon might have occurred in our subjects,
thus revealing higher hippocampus activation during recognitionof male eyes, possibly due to a higher attribution of similarity ascompared to females. Furthermore, activation in rACC has been
related to self-referential processing [33,34]. Emotion recognition
might depend on self-referential processing since, in order to be
successful, the observer tries to covertly mimic the other person’smental state, which then results in shared mental states between
observer and observed person [34]. Together with pSTS andanterior insula activation during emotion vs. gender recognition(regardless of stimulus type), this network might aid in inferring
mental states, and thus predicting other people’s beliefs andintentions by drawing inferences from one’s own and other’s pastand present social information [35].
By contrast, from the regions that showed heightened activation
while inferring mental states vs. gender from female as comparedto male eyes, the precuneus has been repeatedly shown in studies
on theory of mind but also in studies on the self, whereas thedlPFC has been reported during self-regulation [33,34,36].
Furthermore, the dlPFC and the precuneus have shown strong
interconnections [24], and have been related to episodic memory[36,37]. Therefore, their enhanced co-activation during emotion
recognition of female eyes in our male subjects might indicate anincreased attempt for memory retrieval from previous encounters

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stimuli [2]. AVP is an evolutionarily highly conserved mediator in
the regulation of complex social cognition and behavior. Most
brain imaging studies support the view that the effects of oxytocin
and, with less evidence, AVP on social processing and particularly
the emotional component of empathy are mediated by limbic
circuitry, with the amygdala as a core structure [39]. Thus
differences in emotion recognition accuracy to same- vs. oppositesex eyes might be associated with vasopressin modulated changes
in the amygdala.
Mentalizing allows people to empathize with others in order topredict their beliefs, intentions or behavior, or even to deceive
others when needed [33]. Thus, our findings highlight thefunctional significance of the amygdala, particularly for theaffective components of ToM and empathy, and reveal further
evidence that men are less able to infer mental states expressed by
women, which may be accompanied by an amygdala-related
deficit to affectively empathize with women. The finding that men
are superior in recognizing emotions/mental states of other men,as compared to women, might be surprising. From an evolutionary
point of view, accurate interpretations of other men’s rather than
women’s thoughts and intentions, especially threatening cues (alsorelated to amygdala responsiveness [40]), may have been a factor
contributing to survival in ancient times. As men were more
involved in hunting and territory fights, it would have been
important for them to be able to predict and foresee the intentions
and actions of their male rivals.
In further support of this notion, it has been found that, whencompared to women, men are better at identifying facial

expressions of anger [41], while women are more superior inrecognizing fear and sadness [41,42]. Consequently, exposure toangry male as opposed to angry female faces has been shown tolead to enhanced physiological arousal [43] as well as activation of
the ACC [44] in men but not in women.
Since the neural basis of the mentalizing network has also beenfound to be active in studies on self-referential processing, as wellas episodic memory, it is most likely that the ‘core function of thisnetwork is to draw inferences on self- as well as other-related social
information in the past, present and future. This simulation
enables sharing the other’s state based upon one’s own previous
experience and knowledge’ [35]. In evolutionary terms, it also
makes sense that these mentalizing processes operate best in
situations when the other is most similar to oneself.

Acknowledgments
We thank N. Leygraf, and M. Forsting for administrative support and A.
Wormit for assistance with fMRI scanning and data collection. This
research was funded by the Landschaftsverband Rheinland, Germany (Dr.
Schiffer). The funders had no role in study design, data collection andanalysis, decision to publish, or preparation of the manuscript.

Author Contributions
Conceived and designed the experiments: BS BWM. Performed theexperiments: BS CP ERG. Analyzed the data: BS CP HW. Contributed
reagents/materials/analysis tools: BS BWM ERG. Wrote the paper: BS
CP HW.

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