Elements

R E S E A R CH A R T I C L E

Social cognition in the blind brain: A coordinate-based meta-analysis

Maria Arioli1 | Emiliano Ricciardi2 | Zaira Cattaneo1,3

1Department of Psychology, University of

Milano-Bicocca, Milan, Italy

2IMT School for Advanced Studies Lucca,

Lucca, Italy

3IRCCS Mondino Foundation, Pavia, Italy

Correspondence

Zaira Cattaneo, Department of Psychology,

University of Milano-Bicocca, Milan, Italy.

Email: [email protected]

Funding information

Mondino Foundation “Ricerca Corrente”; Italian Ministry of Education University and

Research, Grant/Award Number: 201755TKFE

Abstract

Social cognition skills are typically acquired on the basis of visual information

(e.g., the observation of gaze, facial expressions, gestures). In light of this, a critical

issue is whether and how the lack of visual experience affects neurocognitive mecha-

nisms underlying social skills. This issue has been largely neglected in the literature

on blindness, despite difficulties in social interactions may be particular salient in the

life of blind individuals (especially children). Here we provide a meta-analysis of neu-

roimaging studies reporting brain activations associated to the representation of self

and others' in early blind individuals and in sighted controls. Our results indicate that

early blindness does not critically impact on the development of the “social brain,”

with social tasks performed on the basis of auditory or tactile information driving

consistent activations in nodes of the action observation network, typically active

during actual observation of others in sighted individuals. Interestingly though, acti-

vations along this network appeared more left-lateralized in the blind than in sighted

participants. These results may have important implications for the development of

specific training programs to improve social skills in blind children and young adults.

K E YWORD S

action observation network, activation likelihood estimation, blind, functional magnetic

resonance imaging, meta-analysis, social cognition

1 | INTRODUCTION

Social cognition refers to a complex set of neurocognitive processes

underlying the individuals' ability to decode others' mind to plan

actions in the social environment (Arioli, Crespi, & Canessa, 2018;

Todorov, Harris, & Fiske, 2006). The ability to perceive social informa-

tion (e.g., face expression, posture, voice, action goal) and draw infer-

ences on others' mental states is crucial for survival, cooperation in

social communities, communication and culture (Dolan, 2002). A fun-

damental prerequisite of social cognition is the ability to differentiate

between objects (whose movement is completely explained by physi-

cal forces) and human beings (whose behavior is characterized by

motivations, emotions and believes, which make their actions not

completely predictable) (Fiske & Taylor, 2013; Vogeley, 2017). Social

stimuli, thus, seem to represent a qualitatively different perceptual cate-

gory, mediated by dedicated neurocognitive mechanisms (Maurer,

Grand, & Mondloch, 2002). The prototypical example in this view is the

processing of human faces (Said, Haxby, & Todorov, 2011) and human

bodies in dedicated neural circuitries (i.e., the occipital face area and the

extrastriate body area in the occipitotemporal cortex, and the fusiform

face area and the fusiform body area in the fusiform gyrus; see Bern-

stein, Erez, Blank, & Yovel, 2018; Peelen & Downing, 2007).

The category of social stimuli potentially includes any kind of

information concerning social entities, behavior and words referring

Received: 29 June 2020 Revised: 5 October 2020 Accepted: 31 October 2020

DOI: 10.1002/hbm.25289

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium,

provided the original work is properly cited.

© 2020 The Authors. Human Brain Mapping published by Wiley Periodicals LLC.

Hum Brain Mapp. 2021;42:1243–1256. wileyonlinelibrary.com/journal/hbm 1243

to them (Arioli, Gianelli, & Canessa, 2020). The visual processing of

social stimuli, and human faces in particular, seems to represent the

richest source of information in everyday social life: eyes represent

the most informative social stimuli (Adams & Nelson, 2016), gaze per-

ception is considered a crucial step for mentalizing (Baron-Cohen,

Jolliffe, Mortimore, & Robertson, 1997) and emotional expressions,

produced by the contraction of facial muscles, provide essential social

information (Todorov, Olivola, Dotsch, & Mende-Siedlecki, 2015).

Body postures and gestures are also critical in conveying emotional

states (for a review, see de Gelder, de Borst, & Watson, 2015) and the

study of body emotional perception, both in healthy individuals and

patients, has gained increasing attention in recent years (e.g., Ferrari,

Ciricugno, Urgesi, & Cattaneo, 2019; Ferrari, Papagno, Todorov, &

Cattaneo, 2019; Lenzoni et al., 2020; Soria Bauser, Thoma, &

Suchan, 2012; Thoma et al., 2014; Thoma, Soria Bauser, &

Suchan, 2013). Part of the social information conveyed by visual

cues—such as one's emotional state—may also be acquired via non-

visual sensory modalities: for instance, voice intonation and loudness,

which are features of speech, collectively termed as “prosody”, are

important cues in understanding the speaker's emotional state (see

Mitchell, Elliott, Barry, Cruttenden, & Woodruff, 2003). Nonetheless,

other cues important for social interactions (such as social gestures)

can only be perceived visually. Not surprising, most of the available lit-

erature in social neuroscience has focused on the neural processing of

visual social stimuli. Two brain systems, the action observation/mirror

system and the theory of mind/mentalizing system, are responsible

for the perception and representation of other individuals' states,

actions and intentions (Van Overwalle & Baetens, 2009). Neuroimag-

ing studies identify fronto-parietal and occipito-temporal regions col-

lectively termed the action observation network (AON) as critically

involved in processing others' actions and the meaning underlying

them, via automatic simulation routines (Caspers, Zilles, Laird, &

Eickhoff, 2010). This system allows us to internally, rapidly and intui-

tively simulate observed actions within our own sensorimotor system,

providing an enriched “understanding” of another person's goals and

intentions, on the basis of low-level behavioral input (Caspers

et al., 2010; Iacoboni et al., 2005). In particular, observing actions

recruits the superior and middle temporal gyrus (STG and MTG), infe-

rior parietal lobule (IPL) and inferior frontal gyrus (IFG) (Gardner,

Goulden, & Cross, 2015). In contrast, the mentalizing system allows to

make inferences on others' mental and affective states (Molenberghs,

Johnson, Henry, & Mattingley, 2016), involving the medial precuneus

and temporo parietal junction (TPJ), as well as the ventro-medial and

dorso-medial prefrontal cortex (Amodio & Frith, 2006). Consistent

evidence entails gaze perception as one of the key factors for

mentalizing ability (Calder, Young, Keane, & Dean, 2000). Although

the AON and the mentalizing networks are mainly distinct, they are

likely to play a complementary role during social interactions, and

recent meta-analytic evidence suggests a common involvement of

certain brain regions in both, such as the right pSTS—bordering the

TPJ (Arioli & Canessa, 2019). Particularly, both systems are engaged

during social interactions: the mirror system is responsible for the

preparation of our own actions and the simulation of others' actions,

while the mentalizing system allows to represent others' intentions,

by drawing the capacity to understand the others' thoughts and

beliefs (Sperduti, Guionnet, Fossati, & Nadel, 2014).

Considering the great role of vision in supporting human social

cognition, a critical issue is the impact that blindness, and in particular

the lack of vision since birth, may have on the development of social

cognition, both at the functional and neural level. Blindness may

indeed affect the development of emotional responsiveness and social

skills, possibly predisposing to features of social isolation (including

autism, e.g., Brunes, Hansen, & Heir, 2019; Hobson, Lee, &

Brown, 1999), as well-known by blind educators that are challenged

with the need to develop effective programs to promote social skills

in blind children and youngsters (see Sacks, Kekelis, & Gaylord-

Ross, 1992; Sacks & Wolffe, 2006). Accordingly, studies carried out in

laboratory contexts suggest that blind children show impairment in

several social abilities, such as representing others' mental and affec-

tive states (Brambring & Asbrock, 2010; Dyck, Farrugia, Shochet, &

Holmes-Brown, 2004; Green, Pring, & Swettenham, 2004) and acting

social interactions (Pérez-Pereira & Conti-Ramsden, 2013; Tadic,

Pring, & Dale, 2010). Blind children seem also to exhibit a more limited

repertoire of facial expressions compared to sighted children

(Tröster & Brambring, 1992; see also Webb, 1977). The acquisition of

verbal skills may reduce the impact of early visual deprivation on

social capacities (Bedny & Saxe, 2012), with blind adults being able to

understand other individuals' emotions and mental states in a way

comparable to that of sighted individuals (e.g., Gamond, Vecchi, Fer-

rari, Merabet, & Cattaneo, 2017; Oleszkiewicz, Pisanski, &

Sorokowska, 2017). Nonetheless, it is likely that social cognition may

be mediated by at least partially different strategies and mechanisms

in blind and sighted individuals. For example, blind individuals rely on

partially different strategies in making impressions about social actors

(Ferrari, Vecchi, Merabet, & Cattaneo, 2017), have difficulties in the

posing of emotional expressions (Valente, Theurel, & Gentaz, 2018)

and do not seem to show valence-dependent hemispheric lateraliza-

tion when processing emotions (Gamond et al., 2017). Whereas a con-

sistent body of neuroimaging research has investigated how blindness

affects at the neural level perceptual and cognitive processes, only a

few studies have systematically investigated how blindness affects

the way the brain mediates social processes. Two pioneering fMRI

studies first showed that nonvisual modalities may still drive the

development of the cortical networks underlying action recognition

and Theory of Mind processes (e.g., Bedny, Pascual-Leone, &

Saxe, 2009; Ricciardi et al., 2009). While these observations suggest

that the large-scale functional organization of the “social brain” is

maintained in congenitally blind individuals, there is also evidence that

brain networks specifically devoted to face or voice processing may

develop differently in the absence of visual experience (e.g., Holig,

Focker, Best, Roder, & Buchel, 2014; Pietrini et al., 2004; Van

Ackeren, Barbero, Mattioni, Bottini, & Collignon, 2018).

Meta-analyses are a useful approach that allows to highlight the

consistency of neural pattern across different experimental studies,

by integrating data into a unique statistical analysis. Whereas a recent

meta-analysis by Zhang et al. (2019) has clarified the neurocognitive

1244 ARIOLI ET AL.

mechanisms underlying language, spatial and object processing in

early blind individuals, a comparable approach in the domain of social

cognition has never been implemented. In light of the above, in this

study we implemented a quantitative meta-analysis of the available

neuroimaging literature to provide more solid evidence on: (a) the

brain regions associated with the neural representation of others in

early blind individuals and (b) the specific brain activation in early blind

individuals compared with sighted control individuals. Specifically, we

employed activation likelihood estimation (ALE) in order to draw con-

vergence across neuroimaging experiments on others' representation

in early blind individuals.

2 | MATERIALS AND METHODS

2.1 | Rationale of the meta-analytic approach

We took a quantitative meta-analytic approach to investigate the neu-

ral representation of others in early blind individuals and to unveil

which brain regions are selectively recruited in early blind compared

with sighted control individuals. Critically, with this approach we can

overcome the typical limitations inherent in single neuroimaging stud-

ies, for example, sensitivity to experimental and analytic procedures,

lack of replication studies, as well as small sample size (Carp, 2012).

These constraints are known to increase the likelihood of false nega-

tives (Button et al., 2013), thus pushing researchers toward proce-

dures which, conversely, might promote false positives (Eklund,

Nichols, & Knutsson, 2016; Muller et al., 2018). We thus aimed to

identify the brain regions consistently associated with the neural cod-

ing of others in early blind individuals, over and beyond this process in

sighted control groups. This goal was pursued with ALE, a coordinate-

based meta-analytic approach using the MNI coordinates of peak

locations to summarize and integrate published findings (Turkeltaub,

Eden, Jones, & Zeffiro, 2002). Thus, we ran two separate ALE ana-

lyses: one for blind individuals and one for sighted control individuals.

After that, contrast analyses were conducted between the blind and

the sighted control groups. In particular, we aimed to investigate brain

activations related to others' representation irrespective of the input

sensory modality (i.e., tactile or auditory), the stimulus type

(i.e., vocalizations, sound of actions, words, 3D models of faces to be

tactically explored, etc.), and the specific employed task.

All the inclusion criteria for each dataset were selected by the

first author, and then checked by the other authors. This procedure,

entailing a double check by independent investigators, was aimed to

reduce the chances of a selection bias (Muller et al., 2018).

2.2 | Literature search and study selection: The representation of others in blind and sighted control individuals

We started our survey of the relevant literature by searching for

“early blind fMRI” and “congenitally blind fMRI” on Pubmed (https://

www.ncbi.nlm.nih.gov/pubmed/). The preliminary pool of 1,242 stud-

ies, after duplicates removed, was first screened by title, and then

abstract. We retained only those studies fulfilling the following selec-

tion criteria (see Figure 1 for the detailed study selection process):

1. studies written in English language;

2. empirical fMRI studies, while excluding review and meta-analysis

articles and those employing other techniques, to ensure compara-

ble spatial and temporal resolution;

3. studies reporting whole-brain activation coordinates, rather than

results limited to regions of interest (ROIs) or using small volume

corrected (SVC) analyses. Studies based on ROI or SVC analyses

should be excluded from meta-analyses (Muller et al., 2018),

because a prerequisite is that convergence across experiments is

tested against a null-hypothesis of random spatial associations

across the entire brain under the assumption that each voxel has

the same a priori chance of being activated (Eickhoff, Bzdok, Laird,

Kurth, & Fox, 2012);

4. studies focused on early blind participants, rather than on late

blind participants. In fact, we decided to focus only on early visual

deprivation since neural plasticity phenomena critically depend on

age at blindness onset, with consistent evidence showing that fol-

lowing early visual experience, several brain areas maintain a

vision-dominated response pattern as an outcome of the early

visual experience (e.g., Bedny, Konkle, Pelphrey, Saxe, & Pascual-

Leone, 2010; Voss, Gougoux, Zatorre, Lassonde, & Lepore, 2008;

see also Cattaneo, Vecchi, Monegato, Pece, & Cornoldi, 2007);

5. studies investigating brain activity related to representation of

other individuals (including mental states, physical traits, and

human action recognition, see below for examples) as opposed to

conditions where no representation of other individuals was

involved (such as studies assessing memory capacities, language,

or spatial processing). To this purpose, we selected contrasts

requiring participants to attend to stimuli aimed to elicit a repre-

sentation of other individuals and contrasting this kind of repre-

sentation with baseline conditions where there was no human

representation. The included studies ranged from those requiring

participants to represent others' mental states (Bedny et al., 2009)

to studies comparing responses to human voice processing versus

object sounds (Dormal et al., 2018), to studies comparing haptic

recognition of basic facial expressions versus object discrimination

(Kitada et al., 2013), as well as studies assessing neural activations

in response to sounds produced by human motion (such as foot-

steps, Bedny et al., 2010) or hand-actions (such as cutting paper

with scissors, Ricciardi et al., 2009) compared to non-human envi-

ronmental sounds.

The articles that were excluded, based on titles and abstracts,

included review or meta-analysis studies (31 studies), single case stud-

ies (100 studies), studies not including a blind group (12 studies), stud-

ies assessing other (non-social) cognitive processes (such as spatial

representation) in early blind subjects (137 studies) or in subjects with

cortical visual impairment (34 studies) or in sighted individuals

ARIOLI ET AL. 1245

(646 studies), studies written in non-English language (49 studies) and

studies not employing task-based fMRI technique (185 studies; of

which 11 studies investigated non-social processes in blind subjects,

173 focused on non-social processes in sighted subjects and 1 study

studied social processes in sighted individuals).

From the remaining 48 articles we retained only those fulfilling

the above selection criteria (see Figure 1) and, thus, we excluded

review and meta-analysis articles (11 articles) and studies employing

non-fMRI technique (5 article); studies using ROIs or SVC analyses

(1 study); studies focused on late blind participants (1 study) and stud-

ies that did not focus on others' representation (17 studies).

We included studies fulfilling the above criteria regardless of:

(a) tested sensory modality (e.g., auditory or haptic), (b) experimental

paradigm (e.g., memory or identification tasks). Our aim was indeed to

pool across different experimental paradigms to ensure both general-

izability and consistency of results, within the “others' representation

> non-human representation” comparison inherent in our research

question (Radua & Mataix-Cols, 2012). This selection phase resulted

in 13 studies (out of 374) fulfilling our criteria. In some cases, we

directly contacted the authors to have clarification and more informa-

tion regarding their study.

We then expanded our search for other potentially relevant stud-

ies by carefully examining both the studies quoting, and those quoted

by, each of these papers, alongside a recent meta-analysis on the cog-

nitive processes of blind individuals (Zhang et al., 2019). This second

phase highlighted four further studies fitting our search criteria.

This procedure led to include in the main ALE meta-analysis on

blind participants 17 previously published studies (see Table 1),

resulting from 17 experiments (individual comparisons reported) with

212 subjects and 276 foci. The same procedure led to include 16 pre-

viously published studies, resulting from 16 experiments, with

267 subjects and 331 foci in the ALE analysis regarding the sighted

control group (see Table 1). This number of contrasts is in line with

the recent prescriptions for ALE meta-analyses (Eickhoff et al., 2016;

F IGURE 1 Flowchart of literature search and selection process

1246 ARIOLI ET AL.

TABLE 1 Overview of the 17 studies included in the meta-analysis on the neural representation of other individuals in both blind and sighted groups

N First author/ year Sub Stimuli Task Contrast

Foci

EB

Foci

SC

1 Bedny et al. (2009) 10 EB + 22

SC

Auditory (stories recorded by

female speaker)

True/false question task Mental stories (voice) > noise

control condition; mental

stories > physical stories

11 17

2 Bedny et al. (2010) 10 EB Auditory (sounds) Motion detection task Human sound (footsteps)

> non-human sound

(tones)

5 —

3 Bedny and Saxe (2012) 10 EB + 20

SC

Auditory (words) Semantic judgment task Actions verbs > natural

inanimate objects; mental

verbs > animal nouns

1 8

4 Bedny et al. (2015) 19 EB + 20

SC

Auditory (stories recorded by

female speaker)

“Does this come next?” task

Spoken language > music 6 4

5 Dormal et al. (2018) 16 EB + 15

SC

Auditory (voices) Repetition detection task Voice > scrambled voice;

voice > baseline (silence)

23 27

6 Fairhall et al. (2017) 7 EB + 15 SC Auditory (vocalizations and

statements)

One-back repetition

detection task

Verbal emotional stimuli

(voice) > baseline;

vocalization stimuli >

baseline

57 63

7 Gougoux et al. (2009) 5 EB + 14 SC Auditory (voices) Implicit task (listening) Vocal stimuli > non-vocal

stimuli

3 4

8 Holig et al. (2014) 12 EB + 11

SC

Auditory (voices) Voice identity

recognition

Voice > rest period 90 53

9 Kitada et al. (2013) 17 EB + 22

SC

Haptic (plastic casts) Haptic identification task Face expressions > shoes 3 13

10 Kitada et al. (2014) 24 EB + 28

SC

Haptic (plastic casts) Haptic identification task Hands > nonbiological

control objects

18 23

11 Lewis et al. (2011) 10 EB + 14

SC

Auditory (sounds) Auditory identification

task

Human action sound >

unrecognizable real-world

sounds

5 4

12 Ma and Han (2011) 19 EB + 19

SC

Auditory (statements) Judgment task Others' judgment task >

valence judgment task

2 2

13 Noppeney et al. (2003) 11 EB + 12

SC

Auditory (words) Voice identity

recognition

Hand action words > other

semantic types

1 1

14 Ricciardi et al. (2009) 8 EB + 14 SC Auditory (sounds) Sound recognition Sounds of hand-executed

familiar actions >

environmental sounds

3 6

15 Striem-Amit and

Amedi (2014)

7 EB + 7 SC Auditory (sounds) Visual-to-auditory

sensory-substitution

device (SSD, the

“vOICe”)

Body shape > baseline 12 6

16 van den Hurk et al.

(2017)

14 EB + 18

SC

Auditory (words) One-back task Face and body auditory

conditions > baseline

15 72

17 Hwang and

Matsumoto (2016)

13 EB + 16

SC

Auditory (words) Size judgment task Face parts and celebrity >

places (both famous and

daily)

21 28

Total sub: 212 EB

+ 267 SC

Total foci: 276 331

Note: The majority of these studies employed auditory stimuli (15/17, hence almost 90% of the studies we included), while only two studies used haptic

stimuli (Kitada et al., 2013, 2014).

Abbreviations: EB, early blind individuals; N, progressive study number; SC, sighted control individuals; Sub, subjects.

ARIOLI ET AL. 1247

Muller et al., 2018), and ensures that results would not be driven by

single experiments (see also Zhang et al., 2019).

Importantly, the inclusion of multiple contrasts/experiments from

the same set of subjects can generate dependence across experiment

maps and thus decrease the validity of meta-analytic results. To pre-

vent this issue, we adjusted for within-group effects by pooling the

coordinates from all the relevant contrasts of a study into one experi-

ment (Turkeltaub et al., 2002).

2.3 | Activation likelihood estimation

We performed two distinct ALE analyses, using the GingerALE soft-

ware (Eickhoff et al., 2009), to identify consistently activated regions

associated with the representation of others in both blind and sighted

control groups. We followed the analytic approach previously

described by Arioli and Canessa (2019), based on Eickhoff

et al. (2012). In both meta-analyses, activation foci were initially inter-

preted as the centres of three-dimensional Gaussian probability distri-

butions, to capture the spatial uncertainty associated with each

individual coordinate. All coordinates were reported or converted into

MNI space, using the automatic routine implemented in GingerALE.

The three-dimensional probabilities of all activation foci in a given

experiment were then combined for each voxel, resulting in a modeled

activation (MA) map. The union of these maps produces ALE scores

describing the convergence of results at each brain voxel (Turkeltaub

et al., 2002). To distinguish “true” convergence across studies from

random convergence (i.e., noise), the ALE scores are compared with

an empirically defined null distribution (Eickhoff et al., 2012). The lat-

ter reflects a random spatial association between experiments, with

the within-experiment distribution of foci being treated as a fixed

property. A random-effects inference is thus invoked, by focusing on

the above-chance convergence between different experiments, and

not on the clustering of foci within a specific experiment. From a com-

putational standpoint, deriving this null hypothesis involved sampling

a voxel at random from each MA map, and taking the union of the

resulting values. The ALE score obtained under this assumption of

spatial independence was recorded, and the permutation procedure

iterated 100 times to obtain a sufficient sample of the ALE null distri-

bution. The “true” ALE scores were tested against the ALE scores

obtained under the null distribution and thresholded at p < .001,

corrected for cluster-level family wise error, and the cluster level

threshold was set at p < .05, to identify above-chance convergence in

each analysis (Eickhoff et al., 2012).

The resulting maps were then fed into direct comparisons and

conjunction analyses, within GingerALE, to unveil the common and

specific brain activations between the early blind and sighted control

individuals. A conjunction image was created, using the voxel-wise

minimum value of the included ALE images, to display the similarity

between datasets (Eickhoff et al., 2011). In the same analysis, two

ALE contrast images were created and compared by directly sub-

tracting one input image from the other. To correct for sampling

errors, GingerALE creates such data by pooling the foci in each

dataset and randomly dividing them into two new groupings equiva-

lent in size to the original datasets. An ALE image is created for each

new dataset, then subtracted from the other and compared with the

true data. Permutation calculations are then used to compute a voxel-

wise p value image indicating where the values of the “true data” fall

within the distribution of values in any single voxel. To simplify the

interpretation of ALE contrast images, significant ALE subtraction

scores were converted to Z scores. For between-group contrast ana-

lyses, we used a threshold set at p < .05, corrected for false discovery

rate, and minimum volume size of 100 mm3.

3 | RESULTS

3.1 | Others' representation in early blind individuals

Activations associated with the neural processing of others in early

blind individuals encompassed the regions typically associated with

the AON. These included the posterior portion of the right inferior

frontal gyrus, as well as the inferior and middle temporal cortex,

extending in the right superior temporal sulcus (STS) and the left fusi-

form gyrus (see Table 2 and Figure 2a).

The lack of consistent activation in the parietal cortex, a key node

of the AON, is possibly due to the low number of studies specifically

focusing on hand representation in our database (see Section 4 and

Table 1).

TABLE 2 Neural bases of others' representation in early blind subjects

Cluster # Cluster size (mm3) Brain region x y z

1 872 Right superior temporal gyrus 62 −22 6

2 720 Left fusiform gyrus −52 −58 −4

Left middle temporal gyrus −60 −56 0

3 720 Right inferior frontal gyrus 52 8 28

4 712 Left middle temporal gyrus −54 −70 8

Note: From left to right, the table reports the size (in mm3), stereotaxic coordinates of local maxima and

anatomical labeling of the clusters which were consistently associated with representing others in early

blind subjects.

1248 ARIOLI ET AL.

3.2 | Others' representation in sighted control individuals

Activations associated with representing others in sighted control

groups involved the right superior and middle temporal gyri (see

Table 3 and Figure 2b).

3.3 | Others' representation in early blind and sighted control individuals

A conjunction analysis highlighted no significant common activation

to the processing of other individuals in early blind and sighted control

subjects (Table 4).

The lack of common neural activations between the two

groups during social processing was somehow unexpected and is

probably guided by the low number of studies included. In fact,

the majority of the studies included in our analysis shows over-

lapping activations particularly in the STS, STS/TPJ and the MTG

(see Table 5).

In order to shed light on a possible common neural pattern of

activation in sighted and blind individuals during social tasks, we per-

formed a third meta-analysis with both sighted and early blind individ-

uals (33 experiments included, with 607 foci in 479 subjects). This

additional analysis revealed consistent activation in the right pSTS,

alongside the TPJ, and MTG during the representation of others,

regardless of the group (sighted vs. early blind) (see Table 6 and

Figure 2d).

F IGURE 2 Neural processing of others in early blind (EB) individuals, sighted control (SC) individuals and differences between EB and SC participants. The figure reports the brain structures consistently associated with processing other individuals in EB (a), and SC subjects (b), and the results of direct comparisons and conjunction analysis

between the meta-analyses separately performed on the two different groups (c); this analysis reported no significant common activation to the processing of others in EB and SC, likely due to the low number of studies included in each meta- analysis (17 studies for EB and 16 studies for the SC group). The last panel (d) shows the results of a third meta- analysis carried out considering all studies (both on EB and SC, 33 experiments included) to unveil brain regions consistently engaged during social processing in both groups. All the reported activations survived a statistical threshold of p < .05 corrected for multiple comparisons

ARIOLI ET AL. 1249

3.4 | Others' representation in early blind versus sighted control individuals

In the early blind groups the processing of others was associated with

stronger consistent bilateral activity in left fusiform gyrus and in the

left middle temporal cortex compared to sighted individuals (see

Table 4 and Figure 2c). The reverse comparison highlighted the right

middle/superior temporal gyrus (see Table 4 and Figure 2c).

4 | DISCUSSION

The study of the neural bases of social cognition in the blind brain has

been somehow neglected, with only a few studies specifically investi-

gating whether and how the lack of visual input affects the functional

architecture of the “social brain.” Some studies showed similar pat-

terns of brain activity in early blind and sighted individuals during

tasks tapping on social cognition abilities (Bedny et al., 2009; Ricciardi

et al., 2009), while other studies suggested that social brain networks

develop differently following early visual deprivation (Gougoux

et al., 2009; Holig et al., 2014). These inconsistencies reported in the

neuroimaging literature on social processing in blind individuals may

also reflect possible confounds associated with individual studies, for

example, the influence of experimental and analytic procedures as

well as that of the small sample sizes (Carp, 2012). Moreover, the

effects reported by individual studies are harder to generalize to the

entire target group (here, the early blind), regardless the specific pro-

cedures used (Muller et al., 2018).

In light of this, we pursued a meta-analytic approach to isolate

the most consistent results in the available literature, controlling for

possible confounding effects via stringent criteria for study selection.

In particular, we aimed to investigate: (a) the neural coding of others'

representation in early blind individuals, and (b) the specific brain

regions recruited in early blind compared with sighted control

TABLE 3 Neural bases of others' representation in the sighted control group

Cluster # Cluster size (mm3) Brain region x y z

1 1,032 Right middle temporal gyrus 62 −38 8

Right superior temporal gyrus 56 −32 8

Note: From left to right, the table reports the size (in mm3), stereotaxic coordinates of local maxima and

anatomical labeling of the clusters which were consistently associated with representing others' in

sighted subjects.

TABLE 4 Brain regions showing common and specific activations in the early blind and sighted groups during representation of other individuals

Early blind and sighted

N.A.

Early blind > sighted

Cluster #

Cluster

size (mm3) Brain region x y z

1 616 Left fusiform gyrus −48 −58 −8

Left middle

temporal gyrus

−47 −56 −4

Sighted > early blind

Cluster # Cluster size (mm3) Brain region x y z

1 104 Right middle/superior

temporal gyrus

58 −34 6

Note: From left to right, the table reports the size (in mm3), stereotaxic

coordinates of local maxima and anatomical labeling of the clusters which

were specifically activated in early blind and sighted groups.

TABLE 5 Brain regions showing common activations in early blind and sighted control participants as reported by each study included in the meta-analysis

N First author/year Common activations between SC and EB

1 Bedny et al. (2009) TPJ, PFC and mPFC

2 Bedny et al. (2010) No information provided

3 Bedny and Saxe (2012) MTG

4 Bedny et al. (2015) MTG

5 Dormal et al. (2018) STS

6 Fairhall et al. (2017) MTG, STS, FFA

7 Gougoux et al. (2009) STS

8 Holig et al. (2014) No information provided

9 Kitada et al. (2013) MTG

10 Kitada et al. (2014) MTG, EBA and supramarginal gyrus

11 Lewis et al. (2012) pSTS/MTG

12 Ma and Han (2011) dmPFC and the posterior cingulate

cortex

13 Noppeney et al. (2003) MTG

14 Ricciardi et al. (2009) STG/MTG, premotor cortex,

inferior and middle frontal gyri

(IFG and MFG) and superior and

inferior parietal lobe (SPL, IPL)

15 Striem-Amit and

Amedi (2014)

Multisensory parietal and frontal

areas

16 van den Hurk et al.

(2017)

Ventral-temporal cortex (VTC)

17 Hwang and

Matsumoto (2016)

mPFC, PCC, SFG/MFG, STS/TPJ

and ATL

Note: The majority of the studies reported a common activation in the

superior temporal sulcus (STS), temporo-parietal junction (TPJ) and middle

temporal gyrus (MTG).

1250 ARIOLI ET AL.

individuals, during social processing of others. Although we could

count only on a low number of contrasts, preventing more detailed

analyses such as a direct comparison of nodes of the social brain pos-

sibly differently engaged by auditory and haptic inputs, our sample is

in line with current recommendations for ALE meta-analyses (Eickhoff

et al., 2016; Muller et al., 2018).

Our findings demonstrate that the regions typically associated

with the key nodes of the AON mediate social cognition abilities in

early blind individuals on the basis of non-visual inputs, as they do

during actual observation of others in sighted individuals (Gardner

et al., 2015). In particular, we found consistent overlapping activations

in the middle temporal gyrus bilaterally, alongside the left fusiform

gyrus and the right superior temporal gyrus and finally in the right

inferior frontal gyrus of early blind individuals during others'

processing. The bilateral activation of the middle temporal gyrus has

been previously reported for the AON in a quantitative meta-analysis

on more than 100 studies in sighted individuals (Caspers et al., 2010).

Moreover, similar results, including right frontal activation, are

reported in a meta-analysis investigating brain regions showing mirror

properties through visual and auditory modalities in sighted individ-

uals (Molenberghs, Cunnington, & Mattingley, 2012).

Of note, we found stronger consistent responses in the AON in

the early blind as compared to sighted subjects. This is not surprising,

since the studies we included in the meta-analysis mostly employed

auditory inputs (�90% of the studies, only two studies using haptic

stimuli) that are the typical stimuli on which blind individuals rely on in

social interactions, whereas sighted individuals are mainly guided by

visual cues. This may also account for the different pattern of laterali-

zation observed in the activation of the AON in blind and sighted indi-

viduals, with the former showing more consistent activations in the

left hemisphere particularly in the middle temporal gyrus and fusiform

gyrus, while the reverse comparison highlighted the activation of the

right part of the middle and superior temporal cortex. The different

pattern of hemispheric lateralization may depend on the high familiar-

ity that blind individuals have in recognizing human actions or emo-

tions on the basis of auditory (and haptic) stimuli, with prior evidence

suggesting that action familiarity is associated with increasing activity

in left part of the AON (Gardner et al., 2015; Ricciardi et al., 2009).

An additional analysis carried out on the whole sample of partici-

pants (regardless visual experience) confirmed the common activation

in a right temporal cluster, comprising the STG, the TPJ and the MTG

regions. This cluster appears to be consistently engaged in both

groups during social processing and activations in these regions are

likely to play a more general function in the perception of socially rele-

vant stimuli, which is not bound to visual experience (Fairhall

et al., 2017). These results fit with the involvement of the right STG

and TPJ in a variety of social-cognitive processes (Bahnemann,

Dziobek, Prehn, Wolf, & Heekeren, 2010; Yang, Rosenblau, Keifer, &

Pelphrey, 2015), such as biological motion perception (Beauchamp,

Lee, Haxby, & Martin, 2003; Grossman et al., 2000; Peelen, Wiggett, &

Downing, 2006), mentalizing (Schneider, Slaughter, Becker, &

Dux, 2014; Wolf, Dziobek, & Heekeren, 2010), and emotion attribu-

tion to others on the basis of both visual and auditory/verbal informa-

tion (e.g., Alba-Ferrara, Ellison, & Mitchell, 2012; Ferrari, Schiavi, &

Cattaneo, 2018; Gamond & Cattaneo, 2016; Lettieri et al., 2019;

Redcay, Velnoskey, & Rowe, 2016; Sliwinska & Pitcher, 2018). These

data suggest a two-stage process in which the STS underpins an initial

parsing of a stream of information, whether auditory or visual, into

meaningful discrete elements, whose communicative meaning for

decoding others' behavior and intentions involves more in-depth anal-

ysis associated with increased activation in the TPJ node (Arioli &

Canessa, 2019; Bahnemann et al., 2010; Redcay, 2008). Ethofer

et al. (2006) showed that the pSTS is the input of the prosody

processing system and represents the input to higher-level social cog-

nitive computations, associated with activity in the action observation

system (Gardner et al., 2015), as well as in the mentalizing system

(Schurz, Radua, Aichhorn, Richlan, & Perner, 2014). Accordingly, using

visual stimuli Arioli et al. (2018) pointed to the pSTS as the input for

the social interaction network, which includes key nodes of both

action observation and theory of mind networks. Thus, the STS/TPJ

regions may represent a domain-specific hub associated with the anal-

ysis of the meaning of others' actions, regardless of the stimulation

modality, and highly interconnected with the action observation and

the mentalizing networks.

The lack of activation of the parietal cortex in the early blind, with

the parietal cortex being another key node of the AON in sighted

TABLE 6 Neural bases of others' representation, regardless of visual experience

Cluster # Cluster size (mm3) Brain region x y z

1 6,048 Right superior temporal sulcus/temporo-parietal junction 48 −56 20

Right superior temporal gyrus 64 −22 6

Right superior temporal gyrus 58 −12 2

Right middle temporal gyrus 62 −38 8

Right superior temporal gyrus 56 −32 10

Right superior temporal gyrus 66 −22 −4

Right inferior temporal gyrus 56 −62 −2

Right inferior temporal gyrus 54 −68 2

Right superior temporal sulcus/temporo-parietal junction 58 −50 6

Note: From left to right, the table reports the size (in mm3), stereotaxic coordinates of local maxima and anatomical labeling of the clusters which were

consistently associated with representing others regardless of the group.

ARIOLI ET AL. 1251

individuals, is possibly due to the low number of studies specifically

focused on hand representation in our database (see Pellijeff, Bonilha,

Morgan, McKenzie, & Jackson, 2006; 3/17, see Table 1). Indeed,

Caspers et al. (2010) reported that only observation of hand actions

was consistently associated with activations within parietal cortex,

the observation of non-hand actions was not. Moreover, although

Ricciardi et al. (2009) reported a parietal activation in blind partici-

pants during auditory presentation of hand-executed actions, this acti-

vation was much less extensive (particularly in the superior parietal

cortex) in the blind compared to sighted controls. Another possible

explanation for the lack of parietal activation in the blind in our meta-

analysis (also possibly accounting for the results by Ricciardi

et al., 2009) is that the activation within the parietal part of the AON

may be mainly driven by object-related representations (Caspers

et al., 2010), which are not present in several of the experiments

included in the present analysis. In turn, our studies focused mainly on

voice processing (6/17), and in part on face/body representation

(4/17), this probably being responsible for the consistent activation

we reported in the fusiform gyrus. Indeed, voice processing has been

found to activate the fusiform face area in blind individuals

(i.e., Fairhall et al., 2017).

The findings of this meta-analysis suggest that the AON can

develop despite the lack of any visual experience, with information

acquired in other sensory modalities allowing an efficient representa-

tion of other individuals as agents with specific beliefs and intentions

(vs. objects moved by physical forces). These results may explain why

early blind individuals are able to efficiently interact in the social con-

text and to learn by imitation of others' (e.g., Gamond et al., 2017;

Oleszkiewicz et al., 2017; Ricciardi et al., 2009). Our findings suggest

that regions of the social brain may work on the basis of different sen-

sory inputs, depending on which sensory modality is available. More-

over, our findings are consistent with the results of a recent meta-

analysis by Zhang et al. (2019) and with prior fMRI studies with blind

individuals in the social domain (e.g., Bedny et al., 2009; Ricciardi

et al., 2009) suggesting that brain regions that are consistently rec-

ruited for different functions in sighted individuals, such as the dorsal

fronto-parietal network for spatial function and ventral occipito-

temporal network for object function, and—as shown here—the AON

for social function, maintain their specialization despite the lack of a

normal visual experience. This observation on the “social blind brain”

is in line with the current, more general perspective on the blind brain

that undergoes a functional reorganization due to the lack of visual

experience, but whose large-scale architecture appears to be signifi-

cantly preserved (e.g., Ricciardi, Papale, Cecchetti, & Pietrini, 2020).

Interestingly, we did not find evidence for any cross-modal con-

sistent recruitment of the occipital cortex by social tasks in this meta-

analysis. Cross-modal plasticity typically refers to activation of the

occipital cortex of the early blind in response to input acquired in

other sensory modalities, like hearing and touch (for reviews,

Merabet & Pascual-Leone, 2010; Singh, Phillips, Merabet, &

Sinha, 2018; Voss, 2019), and may account (at least in part) for the

superior perceptual abilities of blind subjects in the spared sensory

modalities (e.g., Battal, Occelli, Bertonati, Falagiarda, &

Collignon, 2020; Bauer et al., 2015). Alternatively, recruitment of the

occipital cortex in the blind has been proposed to also subserve high-

level (cognitive) processing (e.g., Amedi, Raz, Pianka, Malach, &

Zohary, 2003; Bedny, Pascual-Leone, Dodell-Feder, Fedorenko, &

Saxe, 2011; Lane, Kanjlia, Omaki, & Bedny, 2015) suggesting that cor-

tical circuits that are thought to have evolved for visual perception

may come to participate in abstract and symbolic higher-cognitive

functions (see Bedny, 2017). Indeed, recent evidence has shown that

during high-level cognitive tasks (i.e., memory, language and executive

control tasks), there is an increased connectivity between occipital

cortex and associative cortex in the lateral prefrontal, superior parie-

tal, and mid-temporal areas (Abboud & Cohen, 2019), with these

regions being also possibly involved in social perception (Caspers

et al., 2010). In line with this, we would have expected social tasks to

drive activations in the occipital cortex. This was not the case. The

only region that showed a sort of cross-modal response was the fusi-

form face area, in the ventral stream, probably guided by a high num-

ber of studies included in our meta-analysis focusing on voice

processing (cf. Holig et al., 2014; von Kriegstein, Kleinschmidt,

Sterzer, & Giraud, 2005). In this regard, it is also worth noting that

haptic perception by blind individuals of facial expressions and hand

shapes (Kitada et al., 2013, 2014) as well as whole-body shape recog-

nition via a visual-to-auditory sensory substitution device (SSD;

Striem-Amit & Amedi, 2014) led to activations in face and body-

dedicated circuits in the fusiform gyrus, showing that these dedicated

circuits develop even in the absence of a normal visual experience.

Our meta-analysis shows that processes related to representation of

others do not recruit the occipital cortex in the early blind, suggesting

that differently to other cognitive tasks, social tasks may be mediated

by higher-level regions without the need to recruit additional occipital

resources. Even if this might be related to the experimental heteroge-

neities that we highlighted above, the lack of a consistent recruitment

of occipital cortex for social tasks we reported contributes to a better

understanding of the functional role of “visual” areas in the blind

brain.

In conclusion, our findings support the view that the brain of early

blind individuals is functionally organized in the same way of the brain

of sighted individuals although relying on different types of input

(auditory and haptic) (see Bedny et al., 2009; Ricciardi et al., 2009). In

the social domain, this may have important implications for educa-

tional programs for blind children. Early blindness may indeed predis-

pose to features of social isolation, including autism (e.g., Hobson

et al., 1999; Jure, Pogonza, & Rapin, 2016). It is therefore important

to develop training administration guidelines specifically for persons

with visual impairment (Hill-Briggs, Dial, Morere, & Joyce, 2007). In

this regard, an interesting approach would be to develop ad hoc audi-

tory/haptic virtual reality social cognition trainings for children with

blindness or severe visual impairment, as already employed with autis-

tic children and young adults (e.g., Didehbani, Allen, Kandalaft,

Krawczyk, & Chapman, 2016; Kandalaft, Didehbani, Krawczyk, Allen, &

Chapman, 2013). Consistent evidence suggests that audio-based vir-

tual environments may be effective for the transfer of navigation skills

in the blind (Connors, Chrastil, Sanchez, & Merabet, 2014; Sanchez &

1252 ARIOLI ET AL.

Lumbreras, 1999), and haptic virtual perception may be a valid and

effective assistive technology for the education of blind children in

domains like math learning (e.g., Espinosa-Castaneda & Medellin-

Castillo, 2020). This approach—especially audio-based virtual

environments—may thus be extended to the social domain to allow

the safe and non-threatening practice of particular social skills in an

educational setting. In this respect, and considering the importance

for visually impaired children to study in a mainstream school

(e.g., Davis & Hopwood, 2007; Parvin, 2015), school-based social cog-

nitive interventions on the social participation of children with blind-

ness or severe visual impairment would be particularly critical, with

teachers and peers being involved responding and reinforcing blind

children' initiated interactions. A detailed description of the neuro-

cognitive processes underlying social cognition skills in blind individ-

uals is thus critical to tailor training protocols aiming at targeting spe-

cific neuro-cognitive functions. This may have also a translational

clinical impact on the development of non-invasive advanced SSDs

able to translate social cues that are only visually available (such as

face expressions, gestures and body language) to auditory or tactile

feedback that can be processed by the intact social brain of visually

deprived individuals, in terms of more abstract conceptual signals (see

Cecchetti, Kupers, Ptito, Pietrini, & Ricciardi, 2016; Striem-Amit &

Amedi, 2014). These devices may help blind individuals in their inter-

actions both in the physical and virtual (i.e., meetings via Skype, Meet

or others) social world.

ACKNOWLEDGMENTS

This work was supported by a PRIN grant (201755TKFE) to Zaira

Cattaneo and Emiliano Ricciardi by Italian Ministry of Education Uni-

versity and Research, and by Mondino Foundation “Ricerca Corrente”

funds to Zaira Cattaneo. We are grateful to all the authors who kindly

provided unpublished data and/or additional information on their

stud: Marina Bedny, Olivier Collignon, Scott L. Fairhall, Cordula Hölig,

Ryo Kitada, Job van den Hurk and Yanchao Bi.

CONFLICT OF INTEREST

The authors declare no conflict of interest to declare.

DATA AVAILABILITY STATEMENT

Data of this study are available from the corresponding author upon

request.

ORCID

Zaira Cattaneo https://orcid.org/0000-0001-7516-7508

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How to cite this article: Arioli M, Ricciardi E, Cattaneo Z.

Social cognition in the blind brain: A coordinate-based meta-

analysis. Hum Brain Mapp. 2021;42:1243–1256. https://doi.

org/10.1002/hbm.25289

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