Cognition 133 (2014) 9196

Contents lists available at ScienceDirect

Cognition
journal homepage: www.elsevier.com/locate/COGNIT

Brief article

Visual experience inuences the interactions between ngers

and numbers
Virginie Crollen a,b,, Marie-Pascale NoL a, Xavier Seron a, Pierre Mahau a, Franco Lepore b,
Olivier Collignon c,
a

Institut de Recherche en Sciences Psychologiques (IPSY), Centre de Neuroscience Systme et Cognition (NeuroCS), Universit Catholique de Louvain, Belgium
Centre de Recherche en Neuropsychologie et Cognition (CERNEC), Universit de Montral, Canada
c
Centre for Mind/Brain Science, University of Trento, Italy
b

a r t i c l e

i n f o

Article history:
Received 16 September 2013
Revised 13 May 2014
Accepted 7 June 2014
Available online 2 July 2014
Keywords:
Finger-counting
Blindness
Grounded cognition

a b s t r a c t
Though a clear interaction between nger and number representations has been demonstrated, what drives the development of this intertwining remains unclear. Here we tested
early blind, late blind and sighted control participants in two counting tasks, each performed under three different conditions: a resting condition, a condition requiring hands
movements and a condition requiring feet movements. In the resting condition, every
sighted and late blind spontaneously used their ngers, while the majority of early blind
did not. Sighted controls and late blind were moreover selectively disrupted by the interfering hand condition, while the early blind who did not use the nger-counting strategy
remained unaffected by the interference conditions. These results therefore demonstrate
that visual experience plays an important role in implementing the sensori-motor habits
that drive the development of ngernumber interactions.
2014 Elsevier B.V. All rights reserved.

1. Introduction
The nger-based representation of numbers has often
been advocated as an instance of grounded cognition
(e.g., Fischer & Brugger, 2011; Wilson 2002). Since performance on nger discrimination tasks was shown to be a
good predictor of arithmetic abilities (Fayol, Barrouillet, &
Marinthe, 1998; Nol, 2005), it has indeed been argued
that ngers may be the missing tool (Andres, Di Luca, &
Pesenti, 2008) that sustains the assimilation of basic
Corresponding authors. Address: Institut de Recherche en Sciences
Psychologiques (IPSY), Centre de Neuroscience Systme et Cognition,
Universit Catholique de Louvain, Place Cardinal Mercier 10, B-1348
Louvain-la-Neuve, Belgium. Tel.: +32 10 47 40 89 (V. Crollen). Address:
CIMeC Center for Mind/Brain Sciences, University of Trento, via delle
Regole 101, Mattarello, TN, Italy. Tel.: +39 0461 282778; fax: +39 0461
883066 (O. Collignon).
E-mail addresses: virginie.crollen@uclouvain.be (V. Crollen), olivier.
collignon@unitn.it (O. Collignon).
http://dx.doi.org/10.1016/j.cognition.2014.06.002
0010-0277/ 2014 Elsevier B.V. All rights reserved.

numerical abilities or the missing link (Fayol & Seron,

2005) that permits the connection between non-symbolic
numerosities and symbolic arithmetic. Developmental
(Butterworth, 1999a; Costa et al., 2011), neuroimaging
(Harrington et al., 2000; Piazza, Izard, Pinel, Le Bihan, &
Dehaene, 2004; Tschentscher, Hauk, Fischer, &
Pulvermller, 2012), and neuropsychological (Barnes,
Smith-Chant, & Landry, 2005; Gerstmann, 1930;
Thevenot et al., 2014) evidence demonstrating the close
intertwining between ngers and symbolic numbers have
accordingly been accumulated over the last two decades.
Recently, however, it has been highlighted that blind
children used the nger-counting strategy less spontaneously than their sighted peers despite achieving similar
level of counting and nger gnosis (i.e., nger recognition
and localization) performance (Crollen, Mahe, Collignon,
& Seron, 2011). This study has far-reaching implications
since it presumes that the development of ngernumber
interactions (i.e., the associations between symbolic

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V. Crollen et al. / Cognition 133 (2014) 9196

numerical processing and nger movements) relies on sensori-motor habits that are driven by vision. In this paper,
we examined the impact of hand interference on the
counting performance of blind adults. This experiment will
therefore allow us to exclude the idea that nger-counting
develops later in blind people on the basis of non-visual
cues (e.g., kinematic/proprioceptive). It will also allow us
to exclude the idea that nger-counting was present in
blind children but that it did not manifest by an explicit
motor behavior (e.g., absence of voluntary motor activity
but increased cortico-spinal activity of hand muscles;
Andres, Seron, & Olivier, 2007). If nger and number representations actually share common cognitive and/or brain
resources, a motor interference task involving the ngers
should disrupt counting abilities by adding noise in the
shared system.
In the present research, early blind (EB), late blind (LB)
and sighted control adults (SC) were tested with 2 counting tasks and 1 memory task carried out under 3 different
conditions: (1) a control resting condition; (2) a condition requiring the realization of hand movements unrelated to nger-counting; and (3) a condition requiring
the realization of feet movements. If early vision does
not shape the interaction between ngers and the symbolic representation of numbers, all participants should
spontaneously use their ngers to count and should manifest a hand interference effect (i.e., the hand interfering
condition should be more disrupting than the feet condition). In contrast, if early vision is important for the
development of the ngernumber interactions, early
blind individuals should less use their ngers and the
hand interfering condition should not be more disrupting
than the feet condition in this population. Moreover, as
participants were also involved in a working memory task
(listening span test) under the same control and sensorimotor interference conditions, our experiment allowed us
to test whether hand interference effects (Imbo,
Vandierendonck, & Fias, 2011; Michaux, Masson, Pesenti,
& Andres, 2013) would disrupt participants counting performance more than their performance in the listening
span test.

2. Method
2.1. Participants
One group of 15 sighted and two groups of blind participants (11 early and 14 late blinds) took part in the study
(see supplemental Table 1 for a detailed description of
the different groups). In terms of age, the SC did not statistically differ from the EB (p > .2) and LB (p > .1) groups.
Unlike the EB, all LB participants had experienced functional vision before sight loss. At the time of testing, the
participants in both blind groups were totally blind or
had, at the utmost, only rudimentary sensitivity for brightness differences and no patterned vision. In all cases, blindness was attributed to peripheral decits with no
additional neurological problems. Procedures were
approved by the Research Ethics Boards of the University
of Montreal. Experiments were undertaken with the

understanding and written consent of each participant.

Sighted participants were blindfolded when performing
the tasks.
2.2. Conditions
Each of the three tasks (see the tasks section below) was
performed in three different conditions. In a control condition, participants were required to perform the tasks without any constrain. In the hand interference condition,
participants had to perform the tasks while pressing a ball
placed in each hand. Finally, in the foot interference condition, participants had to perform the tasks while pressing a
ball placed beyond each foot.
The rhythm of the interference movements was irregular (between 1500 and 2400 ms) and imposed by a vibrotactile bracelet which was carried on the wrist in the hand
interference condition and on the ankle in the foot interference condition (see supplemental data for a detailed
description of the bracelet).
Before the realization of the experimental tasks, a
5-min training session was performed with the bracelet alone so that participants could train themselves on
the movements. During the experimental tasks, the tactile
stimulations stopped as soon as participants reported the
completion of one trial and started as soon as a new trial
was initiated.
2.3. Tasks
2.3.1. Enumeration task
In order to test the ability to keep track of a number of
enumerated items, participants were required to name a
specic number of exemplars from 10 different target categories (e.g., can you give me 9 names of boys). The target
number ranged from 5 to 9. Three lists of items were created and counterbalanced across participants and conditions. Within a list, each target number was repeated
twice, once in a semantic condition (e.g., can you give me
7 names of tools) and once in a phonological condition
(e.g. can you give me 7 words which begin with the letter
O). Four training trials were presented before the experimental ones. During the instructions, experimenter
emphasized that participants had to stop the enumeration
process (by saying STOP) as soon as they thought
achieved the required target number of words. Participants
were instructed to emphasize accuracy over response
speed. Experimenter noted the number of words uttered
by the participants. As the three lists of stimuli involved
different reaction times in the baseline condition of the
task, only accuracy scores (i.e., number of trials correctly
completed maximum score of 10) were analyzed for each
participant in each condition.
2.3.2. Ordered series manipulation task
In order to test participants ability to count a particular
number of items, participants were asked 15 questions
requiring the manipulation of the letters of the alphabet
(e.g., how many letters are there between c and h?)
and 15 questions requiring the manipulation of the months

V. Crollen et al. / Cognition 133 (2014) 9196

of the year (e.g., how many months are there between

March and September?).
The questions were presented randomly. The same list
of 30 questions was used in the three different conditions.
The target responses were comprised between 5 and 9
and repeated three times with the letters and three times
with the months of the year. Four training trials were presented before the experimental trials. Accuracy scores (i.e.
number of trials correctly completed maximum score of
30) and reaction times were collected for each participant
in each condition. Timing began when the stimulus was
presented and ended when participants gave their
response.
2.3.3. Listening span task
In order to test participants ability to use their working memory, an auditory adaptation of the French version
(Desmette, Hupet, Van der Linden, & Schelstraete, 1995)
of the reading span test (Daneman & Carpenter, 1980)
was presented. This task was used as a control task to
make sure that the potential differences observed in the
other tasks were not due to differences in working
memory. Participants had to listen to a set of recorded
sentences (from 2 to 7) and were instructed to recall
the last word of all the sentences presented in the set.
The task comprised a set of 2 training sentences and 27
experimental sentences (one set of 2 sentences, one set
of 3 sentences, and so on up to the set of 7 sentences).
The inter-sentences interval was 1000 ms. Each trial
started and ended with a 500 ms pink noise. Participants
were required to give their answers after the second
warning tone was emitted. Three lists of sentences were
created and counterbalanced across participants and
conditions. The number of words correctly recalled was
calculated for each participant in each condition (maximum score of 27).
2.4. Procedure
The completion of the experimental procedure
involved two one-hour testing sessions (realized approximately in a week of interval). The control condition was
always performed rst in order to examine whether participants would spontaneously use their ngers to perform the tasks. Order of the two interference conditions
as well as order of the tasks was counterbalanced across
participants.

3. Results
While all SC and LB participants spontaneously used
their ngers to complete the control and foot conditions
of the enumeration and ordered series manipulation tasks,
only 4 EB did so (see supplemental videos 1 and 2). A Chisquared test demonstrated that the EB distribution into
nger-counter and non-nger counter was signicantly
different from the distribution observed in the SC and LB
groups, ps < .001. Two subgroups of EB were therefore
identied: EB who never used their ngers (EB ) and EB

93

who always used their ngers (EB+) (see supplemental

Table 1).
3.1. Enumeration task
Accuracy scores were submitted to a 3 (conditions: control, hand and foot interference)  4 (groups: EB , EB+, LB
and SC)1 ANOVA with repeated measures on the rst factor.
The group effect was not signicant, F(3, 36) = 1.17, p > .3,
g2 = .09. There was, in contrast, a signicant effect of condition, F(2, 72) = 17.67, p < .001, g2 = .33, which was modulated by a condition  group interaction, F(6, 72) = 2.59,
p < .05, g2 = .18. The condition effect was signicant in the
SC group, F(2, 28) = 12.36, p < .001, g2 = .47: accuracy scores
were lower in the hand interference condition by comparison to the control and foot interference conditions. The same
data were observed in the LB group, F(2, 26) = 15.41,
p < .001, g2 = .54, as well as in the EB+ group2, F(2,
6) = 7.87, p < .05, g2 = .72. By contrast, in the EB group2,
participants performed similarly in the three conditions of
the task, F(2, 12) = 0.06, p > .9, g2 = .01 (see Fig. 1).
3.2. Ordered series manipulation task
In order to obtain a general index of performance that
discounts possible criterion shift or speed/accuracy
tradeoff effects, response speed and accuracy were
combined into inverse efciency scores (IES: response
times (RT)/correct response rates; Townsend & Ashby,
1978). As for RT, the lower the score, the better the
performance.
A 3 (conditions: control, hand interference and foot
interference)  4 (groups: EB , EB+, LB and SC)1 repeated
measures ANOVA with group as the between-subject factor and condition as the within subject factor was carried
out on the IES measure. We rst observed a main effect
of condition, F(2, 66) = 13.62, p < .001, g2 = .29. Importantly, we also witnessed a signicant condition  group
interaction, F(6, 66) = 3.94, p < .01, g2 = .26, revealing that
in SC and LB, the hand-interference condition had a particularly negative impact on performance (when compared to
the control condition) while in EB and EB+ the performance was identical in every condition (see Fig. 2). The
EB+ seem however to be more disturbed by the hand
movements than the EB 2.
1
The analyses were also performed on 3 groups (EB, LB, SC) instead of 4
(EB+, EB , LB, SC). In this case, no hand interference effect was observed in
the EB group.
2
As our groups of EB and EB+ were quite small, we also applied the
method recently described by Masson (2011) to compute the posterior
probabilities for H0 and H1. In the ordered manipulation task, this analysis
indicated that the posterior probabilities were .81 for H0 (i.e., the null
hypothesis has 81% chance of being true) and .19 for H1 in the EB group.
According to Rafterys (1995) classication of evidence into weak (.50
.75), positive (.75.95), strong (.95.99), and very strong (>.99), the
probability values obtained for this group therefore provide positive
support for H0 hypothesis. In the EB+ group, the posterior probabilities
were .60 for H0 and .40 for H1 thus providing only weak support for H0. In
the enumeration task, the posterior probabilities in the EB- group were .93
for H0 and .07 for H1. In the EB+ group, the posterior probabilities were .04
for H0 and .96 for H1. These analyses therefore support the idea that the
EB+ are more disturbed by the hand interference condition than the EB .

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V. Crollen et al. / Cognition 133 (2014) 9196

Fig. 1. Results of the enumeration task (maximum score = 10). Error bars denote standard error of the mean. EB
ngers; EB+, the early blind who used their ngers; LB, the late blind and SC, the sighted controls.

Fig. 2. Results of the ordered series manipulation task. Error bars denote standard error of the mean. EB
EB+, the early blind who used their ngers; LB, the late blind and SC, the sighted controls.

3.3. Listening span task

A repeated measures ANOVA with condition (control,
hand interference, foot interference) as the within-subject
factor and group (EB+, EB , LB, SC)1 as the between subject
factor was run on the accuracy scores. Results only showed
a marginally signicant effect of group, F(3, 36) = 2.42,
p = .08, g2 = .17. EB (M = 24.76 0.83) performed better
than LB (M = 22.33 0.58) and SC (M = 22.67 0.57). No
other difference was signicant (see Fig. 3).

4. Discussion
The study of blind individuals offers the unique opportunity to examine how visual experience shapes cognition in
the context of extreme changes in the environmental input
(Bedny & Saxe, 2012; Crollen, Dormal, Seron, Lepore, &
Collignon, 2013). Here, we studied visually deprived individuals in order to obtain new insights into the origins of
the interactions between ngers and symbolic numbers.

are the early blind who did not use their

are the early blind who did not use their ngers;

While all SC and LB participants used their ngers to

perform the counting tasks, the majority of EB did not.
Moreover, hand movements interfered with counting in
SC and LB but not in the EB who did not use their ngers
in the enumeration tasks. All together, these results suggest that developmental vision is instrumental in implementing the close connection between ngers and
counting. However, since a minority of EB uses their ngers to count and shows specic manual interferences
(EB+), blindness does not seem to, by itself, skim off ngernumber interactions.
Interestingly, all EB (EB and EB+) stated that they
never learned nger-counting at school or with their parents. So, we do not know why four of the EB used a nger
counting strategy. It is possible that non-visual cues (kinematic/proprioceptive) are employed to develop the nger
counting habit. Several classes of afferent signals from
the periphery may indeed provide information about the
location of the limbs, including receptors in joints signaling
exion or extension, from the skin signaling stretch, and
from muscle spindles signaling contraction or lengthening

V. Crollen et al. / Cognition 133 (2014) 9196

Fig. 3. Results of the listening span test (maximum score = 27). Error bars denote standard error of the mean. EB
ngers; EB+, the early blind who used their ngers; LB, the late blind and SC, the sighted controls.

95

are the early blind who did not use their

(Proske & Gandevia, 2012). However, the fact that the

majority of EB do not spontaneously use their ngers and
do not suffer from specic hand interference suggests that
these cues are less efcient than the visual ones to implement nger-counting strategies. Vision thus probably provides an important but not mandatory interface to confer
to ngers a useful value as a tool to support counting.
On another hand, the fact that EB realize the task with
equal performance as the 3 other groups suggests that the
development of the symbolic numerical system is exible
enough to rely on different kinds of sensory and cognitive
strategies. Among the hypotheses that still need to be
tested, it may be presumed that EB make a more appropriate use of working memory capacities (Castronovo &
Delvenne, 2013; Crollen & Mahe et al., 2011; Crollen,
Seron, & Nol, 2011; Withagen, Kappers, Vervloed,
Knoors, & Verhoeven, 2013).
In the literature, several brain mapping studies suggest
that there is a shared neural network for number and nger processing, including the parietal areas, the precentral
gyrus and the primary motor cortex (Andres, Michaux, &
Pesenti, 2012; Andres et al., 2007; Harrington et al.,
2000; Piazza et al., 2004; Tschentscher et al., 2012; Zago
et al., 2001)3. Two prevailing views have been recently
debated in order to explain the origin of this neuro-anatomical overlap: the functionalist and the redeployment hypotheses. According to the functionalist view, neuronal
activations for number processing and nger movements
are correlated in adulthood because ngers are used by children while learning counting and basic arithmetic operations (Butterworth, 1999b). The redeployment view
assumes that functional circuits originally evolved for nger
representation have since been redeployed to support the
representation of number and now serves both uses
(Penner-Wilger & Anderson, 2008; Penner-Wilger &
Anderson, 2013). For the functionalist theory, re-use

happens over the course of development whereas it happens

over the course of evolution for the redeployment hypothesis (Anderson, 2010; Penner-Wilger & Anderson, 2013). One
key prediction of the redeployment hypothesis is therefore
that individuals with intact nger gnosis who did not use
their ngers to represent quantities during development
should nevertheless show activation in the nger circuit
during tasks requiring the representation of numbers
(Penner-Wilger & Anderson, 2013). Our observation that
EB are not impaired in the interfering hand condition
(which should induce noise in the pre-existing overlapping
circuits for number and nger processing) compellingly
argues against the redeployment view. We therefore suggest
that vision provides an ideal interface to trigger the development of ngernumber interactions. In the absence of
vision, the development of this association is less likely,
and other sensory/cognitive strategies are used to support
counting. Our prediction is that EB would not show overlapping brain circuitry representing ngers and counting. It
could therefore be highly interesting to investigate how
the well-known crossmodal reorganization of the occipitoparietal network in early blind individuals (Collignon,
Davare, Olivier, & De Volder, 2009; Collignon, Voss,
Lassonde, & Lepore, 2009; Collignon et al., 2011; Dormal,
Lepore, & Collignon, 2012) affects the circuitry representing
space, number and nger processing.
In summary, our study provides some breakthroughs in
our understanding of the relation between ngers and
counting by demonstrating that: (1) the use of ngers is
not mandatory to achieve optimal performance in counting, (2) vision plays an important role in the establishment
of ngernumber interactions, probably because it provides an ideal platform to relieve the memory load during
counting; (3) the development of this intertwining
depends on experience and is not the product of and inherited redeployment of function.

3
While some studies did not involve any explicit nger movements
(Tschentscher et al., 2012), many of the above mentioned research used
tasks which are spatial in nature by involving movements of ngers
(Harrington et al., 2000), pointing/grasping (Simon, Mangin, Cohen, Le
Bihan, & Dehaene, 2002), or mapping nger locations to a spatial position
(Andres et al., 2012). It is therefore difcult to strongly argue that the
parietal cortex is involved in nger representation per s.

Acknowledgments
Authors wish to thank Giulia Dormal for her help in preliminary data collection and Samuel Di Luca for his help in
editing the videos. This research and the authors were supported by the Canada Research Chair Program (FL), the

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V. Crollen et al. / Cognition 133 (2014) 9196

Canadian Institutes of Health Research (FL), the Fondation

Sainte-Justine (OC), the Belgian National Funds for
Scientic Research (VC), the Fonds Spciaux de Recherche
of the Universit Catholique de Louvain (FSR; VC) and a
WBI. World grant (VC). European Research Council ERC2013-StG Project no. 337573 MADVIS attributed to OC.
Appendix A. Supplementary material
Supplementary data associated with this article can be
found, in the online version, at http://dx.doi.org/10.1016/
j.cognition.2014.06.002.
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