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Cognition
journal homepage: www.elsevier.com/locate/COGNIT
Department of Brain and Cognitive Sciences, Massachusetts Institute of Technology, Cambridge, MA 02139, USA
Department of Psychological and Brain Sciences, Johns Hopkins University, Baltimore, MD 21218, USA
a r t i c l e
i n f o
Article history:
Received 13 February 2013
Revised 19 February 2014
Accepted 13 April 2014
Keywords:
Blindness
Theory of mind
Experience
Representation
fMRI
Multivoxel pattern analysis (MVPA)
a b s t r a c t
Blind peoples inferences about how other people see provide a window into fundamental
questions about the human capacity to think about one anothers thoughts. By working
with blind individuals, we can ask both what kinds of representations people form about
others minds, and how much these representations depend on the observer having had
similar mental states themselves. Thinking about others mental states depends on a specific group of brain regions, including the right temporo-parietal junction (RTPJ). We investigated the representations of others mental states in these brain regions, using multivoxel
pattern analyses (MVPA). We found that, first, in the RTPJ of sighted adults, the pattern of
neural response distinguished the source of the mental state (did the protagonist see or
hear something?) but not the valence (did the protagonist feel good or bad?). Second, these
neural representations were preserved in congenitally blind adults. These results suggest
that the temporo-parietal junction contains explicit, abstract representations of features
of others mental states, including the perceptual source. The persistence of these representations in congenitally blind adults, who have no first-person experience with sight,
provides evidence that these representations emerge even in the absence of relevant
first-person perceptual experiences.
2014 Elsevier B.V. All rights reserved.
1. Introduction
Imagine a friend tells you that last night, looking out the
window onto a dark, rainy street, she saw her boyfriend get
into a car with a strange woman, and drive away. Your
reaction will depend on many inferences about her
thoughts and feelings. You will recognize that she believes
her boyfriend is being unfaithful, and feels betrayed. You
might also note the source of her belief, and question
how clearly she could see at a distance and in the dark.
Perhaps she was mistaken about the identity of the man
getting into the car, or about the driver; maybe the driver
Corresponding author. Address: 77 Massachusetts Ave 46-4021C,
Cambridge, MA 02210, USA. Tel.: +1 857 272 0733.
E-mail address: jorie@mit.edu (J. Koster-Hale).
http://dx.doi.org/10.1016/j.cognition.2014.04.006
0010-0277/ 2014 Elsevier B.V. All rights reserved.
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terns generated by items within a condition are more similar to each other (within-condition correlation) than to
the neural patterns generated by items in the other conditions (across-condition correlation). If we find that
within-condition correlations are reliably higher than
across-condition correlations, we can conclude that there
are reliable neural pattern generated by different items
within a condition, and that these neural patterns are distinct from one condition to another. Together, this suggests
that the region is sensitive to the category distinction
items within a category are coded in a similar way, with
distinguishable codes for different categories.
Here, we conducted within-ROI pattern analyses, independently testing for information about belief source and
valence in the regions identified in the independent functional localizer. To compare seeing and hearing beliefs, we
collapsed across good and bad valence; to compare good
and bad valence, we collapsed across seeing and hearing.
Following Haxby (2001), each participants data were
divided into even and odd runs (partitions) and then the
mean response (beta value) of every voxel in the ROI was
calculated for each condition. Because each participant
read 8 stories about hearing, seeing, feeling good, and feeling bad, each partition contained the average response to 4
individual stories. The pattern of response was the vector
of beta values across voxels within the participants individual ROI. To determine the within-condition correlation,
the pattern in one (e.g. even) partition was correlated with
the pattern for the same condition in the opposite
(e.g. odd) partition; to determine the across-condition
correlations the pattern was compared to the opposite
condition, across partitions (Fig. 2).
For each condition pair (e.g. seeing vs. hearing) in each
individual, an index of classification was calculated as the
within-condition correlation (e.g. the correlation of one
half of the seeing stories to the other half of seeing stories,
averaged with the correlation of one half of the hearing to
the other half of hearing stories) minus the acrosscondition correlation (e.g. the correlation of seeing stories
compared to hearing stories). To allow for direct comparison of correlation coefficients, we transformed all r values
using Fishers Z transform. A region successfully classified a
category of stimuli if, across individuals, the withincondition correlation was higher than the across-condition
correlation, using a Students T complementary cumulative
distribution function.
This procedure implements a simple linear decoder.
Linear decoding, while in principle less flexible and less
powerful than non-linear decoding, is preferable both theoretically and empirically. A non-linear classifier can
decode nearly any arbitrary feature contained implicitly
within an ROI, reflecting properties of the pattern analysis
algorithm rather than the brain, which makes successful
classification largely uninformative (Cox & Savoy, 2003;
DiCarlo & Cox, 2007; Goris & Op de Beeck, 2009;
Kamitani & Tong, 2005; Norman et al., 2006). Moreover,
linear codes have been argued to be a more neurally plausible way of making information available to the next layer
of neurons (Bialek, Rieke, Van Steveninck, & Warland,
1991; Butts et al., 2007; DiCarlo & Cox, 2007; Naselaris,
Kay, Nishimoto, & Gallant, 2011; Rolls & Treves, 2011).
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Fig. 2. Split half MVPA analysis. The data from each condition (e.g.
hearing and seeing) were divided by run, and for each participant, we
asked whether the within condition correlations (blue arrows) were
higher than the across conditions correlations (gray arrows). Data from
one blind participant: within-condition correlation = 1.3; across-condition correlation = 0.8. (For interpretation of the references to color in this
figure legend, the reader is referred to the web version of this article.)
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Williams, 2011), we used difference scores (within condition correlation across condition correlation) as the
dependent variable. We found a main effect of ROI
(F(4,28) = 3.03, p = 0.03, partial g2 = 0.27), suggesting that
the regions contain varying amounts of information about
source modality in their neural pattern.
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Fig. 3. MVPA results. (A) Canonical theory of mind brain regions; for all analyses, theory of mind brain regions were individual defined in each participant.
(B) Sighted adults (n = 13) show pattern discrimination for beliefs based on seeing vs. hearing in the right and left TPJ, but not in any other theory of mind
region. (C) These representations of seeing and hearing persist in the RTPJs of blind adults (n = 9). Error bars show the standard error of within-participant
difference scores (within condition correlation - across condition correlation).
Table 1
Z scores for within versus across condition comparisons of seeing and hearing.
Region
Within
Across
Significance
Sighted
RTPJ
LTPJ
PC
DMPFC
VMPFC
1.1 0.2
0.82 0.2
0.86 0.2
0.7 0.1
0.58 0.2
0.95 0.2
0.64 0.2
0.78 0.2
0.5 0.2
0.47 0.2
Blind
RTPJ
LTPJ
PC
DMPFC
VMPFC
1.1 0.2
1.3 0.3
0.71 0.2
0.83 0.2
0.56 0.2
0.93 0.3
1.3 .03
0.59 0.2
0.73 0.1
0.34 0.2
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support of this project by an NSF Graduate Research Fellowship (#0645960 to JKH) and an NSF CAREER award
(#095518), NIH (1R01 MH096914-01A1), and the Packard
Foundation (to RS).
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