numerical encoding in early visual cortex€¦ · visual visual cortex cortex q1 q1 1 1 q1 q1 0 0...
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Special
Special
Special
Special
Special
Special
Special issue:
issue:
issue:
issue:
issue:
issue:
issue: Research
Research
Research
Research
Research
Research
Research report
report
report
report
report
report
report
Numerical
Numerical
Numerical
Numerical
Numerical
Numerical
NumericalNumerical encoding
encoding
encoding
encoding
encoding
encoding
encodingencoding in
in
in
in
in
in
inin early
early
early
early
early
early
earlyearly visual
visual
visual
visual
visual
visual
visualvisual cortex
cortex
cortex
cortex
cortex
cortex
cortexcortexQ1Q1
Q1
Q1
Q1
Q1
Q11
1
1
1
1
1
1
Q1
Q1
Q1
Q1
Q1
Q1
Q10
0
0
0
0
0
0 Nicholas
Nicholas
Nicholas
Nicholas
Nicholas
Nicholas
Nicholas K.
K.
K.
K.
K.
K.
K. DeWind
DeWind
DeWind
DeWind
DeWind
DeWind
DeWind a
a
a
a
a
a
a,
,
,
,
,
,
,*
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*
*
*
*,
,
,
,
,
,
, Joonkoo
Joonkoo
Joonkoo
Joonkoo
Joonkoo
Joonkoo
Joonkoo Park
Park
Park
Park
Park
Park
Park b
b
b
b
b
b
b,
,
,
,
,
,
, Marty
Marty
Marty
Marty
Marty
Marty
Marty G.
G.
G.
G.
G.
G.
G. Woldorff
Woldorff
Woldorff
Woldorff
Woldorff
Woldorff
Woldorff c
c
c
c
c
c
c and
and
and
and
and
and
andElizabeth
Elizabeth
Elizabeth
Elizabeth
Elizabeth
Elizabeth
Elizabeth M.
M.
M.
M.
M.
M.
M. Brannon
Brannon
Brannon
Brannon
Brannon
Brannon
Brannon a
a
a
a
a
a
a
a
a
a
a
a
a
a Department
Department
Department
Department
Department
Department
Department of
of
of
of
of
of
of Psychology,
Psychology,
Psychology,
Psychology,
Psychology,
Psychology,
Psychology, University
University
University
University
University
University
University of
of
of
of
of
of
of Pennsylvania,
Pennsylvania,
Pennsylvania,
Pennsylvania,
Pennsylvania,
Pennsylvania,
Pennsylvania, Philadelphia,
Philadelphia,
Philadelphia,
Philadelphia,
Philadelphia,
Philadelphia,
Philadelphia, PA,
PA,
PA,
PA,
PA,
PA,
PA, USA
USA
USA
USA
USA
USA
USAb
b
b
b
b
b
b Department
Department
Department
Department
Department
Department
Department of
of
of
of
of
of
of Psychological
Psychological
Psychological
Psychological
Psychological
Psychological
Psychological and
and
and
and
and
and
and Brain
Brain
Brain
Brain
Brain
Brain
Brain Sciences,
Sciences,
Sciences,
Sciences,
Sciences,
Sciences,
Sciences, University
University
University
University
University
University
University of
of
of
of
of
of
of Massachusetts
Massachusetts
Massachusetts
Massachusetts
Massachusetts
Massachusetts
Massachusetts Amherst,
Amherst,
Amherst,
Amherst,
Amherst,
Amherst,
Amherst, Amherst,
Amherst,
Amherst,
Amherst,
Amherst,
Amherst,
Amherst, MA,
MA,
MA,
MA,
MA,
MA,
MA, USA
USA
USA
USA
USA
USA
USAQ1
Q1
Q1
Q1
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Q1c
c
c
c
c
c
c Department
Department
Department
Department
Department
Department
Department of
of
of
of
of
of
of Psychology
Psychology
Psychology
Psychology
Psychology
Psychology
Psychology and
and
and
and
and
and
and Neuroscience,
Neuroscience,
Neuroscience,
Neuroscience,
Neuroscience,
Neuroscience,
Neuroscience, Duke
Duke
Duke
Duke
Duke
Duke
Duke University,
University,
University,
University,
University,
University,
University, Durham,
Durham,
Durham,
Durham,
Durham,
Durham,
Durham, NC,
NC,
NC,
NC,
NC,
NC,
NC, USA
USA
USA
USA
USA
USA
USA
a r t i c l e i n f o
a r t i c l e i n f o
a r t i c l e i n f o
a r t i c l e i n f o
a r t i c l e i n f o
a r t i c l e i n f o
a r t i c l e i n f o
Article
Article
Article
Article
Article
Article
Article history:
history:
history:
history:
history:
history:
history:
Received
Received
Received
Received
Received
Received
Received 17
17
17
17
17
17
17 December
December
December
December
December
December
December 2017
2017
2017
2017
2017
2017
2017
Reviewed
Reviewed
Reviewed
Reviewed
Reviewed
Reviewed
Reviewed 23
23
23
23
23
23
23 January
January
January
January
January
January
January 2018
2018
2018
2018
2018
2018
2018
Revised
Revised
Revised
Revised
Revised
Revised
Revised 13
13
13
13
13
13
13 March
March
March
March
March
March
March 2018
2018
2018
2018
2018
2018
2018
Accepted
Accepted
Accepted
Accepted
Accepted
Accepted
Accepted 27
27
27
27
27
27
27 March
March
March
March
March
March
March 2018
2018
2018
2018
2018
2018
2018
Published
Published
Published
Published
Published
Published
Published online
online
online
online
online
online
online xxx
xxx
xxx
xxx
xxx
xxx
xxx
Keywords:
Keywords:
Keywords:
Keywords:
Keywords:
Keywords:
Keywords:
Numerosity
Numerosity
Numerosity
Numerosity
Numerosity
Numerosity
Numerosity
Intraparietal
Intraparietal
Intraparietal
Intraparietal
Intraparietal
Intraparietal
Intraparietal area
area
area
area
area
area
area
Early
Early
Early
Early
Early
Early
Early visual
visual
visual
visual
visual
visual
visual cortex
cortex
cortex
cortex
cortex
cortex
cortexQ3
Q3
Q3
Q3
Q3
Q3
Q3
a b s t r a c t
a b s t r a c t
a b s t r a c t
a b s t r a c t
a b s t r a c t
a b s t r a c t
a b s t r a c t
The
The
The
The
The
The
The ability
ability
ability
ability
ability
ability
ability to
to
to
to
to
to
to estimate
estimate
estimate
estimate
estimate
estimate
estimate the
the
the
the
the
the
the number
number
number
number
number
number
number in
in
in
in
in
in
in a
a
a
a
a
a
a visual
visual
visual
visual
visual
visual
visual array
array
array
array
array
array
array arose
arose
arose
arose
arose
arose
arose early
early
early
early
early
early
early in
in
in
in
in
in
in evolution,
evolution,
evolution,
evolution,
evolution,
evolution,
evolution, develops
develops
develops
develops
develops
develops
develops early
early
early
early
early
early
early
in
in
in
in
in
in
in hu
hu
hu
hu
hu
hu
human
man
man
man
man
man
man developme
developme
developme
developme
developme
developme
development,
nt,
nt,
nt,
nt,
nt,
nt, and
and
and
and
and
and
and is
is
is
is
is
is
is correla
correla
correla
correla
correla
correla
correlated
ted
ted
ted
ted
ted
ted with
with
with
with
with
with
with mathematical
mathematical
mathematical
mathematical
mathematical
mathematical
mathematical ability.
ability.
ability.
ability.
ability.
ability.
ability. Previous
Previous
Previous
Previous
Previous
Previous
Previous work
work
work
work
work
work
work with
with
with
with
with
with
with
visually
visually
visually
visually
visually
visually
visually presented
presented
presented
presented
presented
presented
presented arrays
arrays
arrays
arrays
arrays
arrays
arrays indicates
indicates
indicates
indicates
indicates
indicates
indicates that
that
that
that
that
that
that the
the
the
the
the
the
the intraparietal
intraparietal
intraparietal
intraparietal
intraparietal
intraparietal
intraparietal sulcus
sulcus
sulcus
sulcus
sulcus
sulcus
sulcus (IPS)
(IPS)
(IPS)
(IPS)
(IPS)
(IPS)
(IPS) represents
represents
represents
represents
represents
represents
represents number.
number.
number.
number.
number.
number.
number.
However,
However,
However,
However,
However,
However,
However, it
it
it
it
it
it
it is
is
is
is
is
is
is not
not
not
not
not
not
not clear
clear
clear
clear
clear
clear
clear if
if
if
if
if
if
if the
the
the
the
the
the
the number
number
number
number
number
number
number signal
signal
signal
signal
signal
signal
signal originates
originates
originates
originates
originates
originates
originates in
in
in
in
in
in
in IPS
IPS
IPS
IPS
IPS
IPS
IPS or
or
or
or
or
or
or is
is
is
is
is
is
is propagated
propagated
propagated
propagated
propagated
propagated
propagated from
from
from
from
from
from
from earlier
earlier
earlier
earlier
earlier
earlier
earlier
visual
visual
visual
visual
visual
visual
visual areas.
areas.
areas.
areas.
areas.
areas.
areas. Previous
Previous
Previous
Previous
Previous
Previous
Previous work
work
work
work
work
work
work from
from
from
from
from
from
from our
our
our
our
our
our
our group
group
group
group
group
group
group has
has
has
has
has
has
has demons
demons
demons
demons
demons
demons
demonstrated
trated
trated
trated
trated
trated
trated a
a
a
a
a
a
a rapidly
rapidly
rapidly
rapidly
rapidly
rapidly
rapidly instantiated
instantiated
instantiated
instantiated
instantiated
instantiated
instantiated rep-
rep-
rep-
rep-
rep-
rep-
rep-
resentation
resentation
resentation
resentation
resentation
resentation
resentation of
of
of
of
of
of
of number
number
number
number
number
number
number in
in
in
in
in
in
in low-level
low-level
low-level
low-level
low-level
low-level
low-level regions
regions
regions
regions
regions
regions
regions of
of
of
of
of
of
of visual
visual
visual
visual
visual
visual
visual cortex
cortex
cortex
cortex
cortex
cortex
cortex using
using
using
using
using
using
using the
the
the
the
the
the
the high
high
high
high
high
high
high temporal
temporal
temporal
temporal
temporal
temporal
temporal reso-
reso-
reso-
reso-
reso-
reso-
reso-
lution
lution
lution
lution
lution
lution
lution of
of
of
of
of
of
of event-related
event-related
event-related
event-related
event-related
event-related
event-related electro-encephalography
electro-encephalography
electro-encephalography
electro-encephalography
electro-encephalography
electro-encephalography
electro-encephalography (EEG
(EEG
(EEG
(EEG
(EEG
(EEG
(EEG).
).
).
).
).
).
). Here,
Here,
Here,
Here,
Here,
Here,
Here, we
we
we
we
we
we
we use
use
use
use
use
use
use a
a
a
a
a
a
a rapid
rapid
rapid
rapid
rapid
rapid
rapid event-related
event-related
event-related
event-related
event-related
event-related
event-related
functional
functional
functional
functional
functional
functional
functional magnetic
magnetic
magnetic
magnetic
magnetic
magnetic
magnetic resonance
resonance
resonance
resonance
resonance
resonance
resonance imaging
imaging
imaging
imaging
imaging
imaging
imaging (fMRI)
(fMRI)
(fMRI)
(fMRI)
(fMRI)
(fMRI)
(fMRI) paradigm
paradigm
paradigm
paradigm
paradigm
paradigm
paradigm and
and
and
and
and
and
and find
find
find
find
find
find
find convergent
convergent
convergent
convergent
convergent
convergent
convergent evidence
evidence
evidence
evidence
evidence
evidence
evidence for
for
for
for
for
for
for
a
a
a
a
a
a
a number
number
number
number
number
number
number signal
signal
signal
signal
signal
signal
signal in
in
in
in
in
in
in low-level
low-level
low-level
low-level
low-level
low-level
low-level visual
visual
visual
visual
visual
visual
visual cortex
cortex
cortex
cortex
cortex
cortex
cortex (areas
(areas
(areas
(areas
(areas
(areas
(areas V1,
V1,
V1,
V1,
V1,
V1,
V1, V2,
V2,
V2,
V2,
V2,
V2,
V2, and
and
and
and
and
and
and V3).
V3).
V3).
V3).
V3).
V3).
V3). Employing
Employing
Employing
Employing
Employing
Employing
Employing a
a
a
a
a
a
a stringent
stringent
stringent
stringent
stringent
stringent
stringent set
set
set
set
set
set
set
of
of
of
of
of
of
of stimulus
stimulus
stimulus
stimulus
stimulus
stimulus
stimulus controls,
controls,
controls,
controls,
controls,
controls,
controls, we
we
we
we
we
we
we demonstrate
demonstrate
demonstrate
demonstrate
demonstrate
demonstrate
demonstrate that
that
that
that
that
that
that this
this
this
this
this
this
this signal
signal
signal
signal
signal
signal
signal cannot
cannot
cannot
cannot
cannot
cannot
cannot be
be
be
be
be
be
be explaine
explaine
explaine
explaine
explaine
explaine
explained
d
d
d
d
d
d by
by
by
by
by
by
by the
the
the
the
the
the
the total
total
total
total
total
total
total
extent
extent
extent
extent
extent
extent
extent of
of
of
of
of
of
of the
the
the
the
the
the
the array,
array,
array,
array,
array,
array,
array, the
the
the
the
the
the
the density
density
density
density
density
density
density of
of
of
of
of
of
of the
the
the
the
the
the
the items
items
items
items
items
items
items in
in
in
in
in
in
in the
the
the
the
the
the
the array,
array,
array,
array,
array,
array,
array, the
the
the
the
the
the
the aggregate
aggregate
aggregate
aggregate
aggregate
aggregate
aggregate visual
visual
visual
visual
visual
visual
visual area
area
area
area
area
area
area of
of
of
of
of
of
of the
the
the
the
the
the
the
items,
items,
items,
items,
items,
items,
items, the
the
the
the
the
the
the size
size
size
size
size
size
size of
of
of
of
of
of
of individual
individual
individual
individual
individual
individual
individual items,
items,
items,
items,
items,
items,
items, the
the
the
the
the
the
the proportion
proportion
proportion
proportion
proportion
proportion
proportion of
of
of
of
of
of
of the
the
the
the
the
the
the array
array
array
array
array
array
array covered
covered
covered
covered
covered
covered
covered by
by
by
by
by
by
by items,
items,
items,
items,
items,
items,
items, nor
nor
nor
nor
nor
nor
nor the
the
the
the
the
the
the
overall
overall
overall
overall
overall
overall
overall scale
scale
scale
scale
scale
scale
scale of
of
of
of
of
of
of the
the
the
the
the
the
the array
array
array
array
array
array
array and
and
and
and
and
and
and items.
items.
items.
items.
items.
items.
items. Our
Our
Our
Our
Our
Our
Our findings
findings
findings
findings
findings
findings
findings thus
thus
thus
thus
thus
thus
thus provide
provide
provide
provide
provide
provide
provide strong
strong
strong
strong
strong
strong
strong support
support
support
support
support
support
support for
for
for
for
for
for
for the
the
the
the
the
the
the hy-
hy-
hy-
hy-
hy-
hy-
hy-
pothesis
pothesis
pothesis
pothesis
pothesis
pothesis
pothesis that
that
that
that
that
that
that number
number
number
number
number
number
number is
is
is
is
is
is
is rapidly
rapidly
rapidly
rapidly
rapidly
rapidly
rapidly and
and
and
and
and
and
and directly
directly
directly
directly
directly
directly
directly encoded
encoded
encoded
encoded
encoded
encoded
encoded early
early
early
early
early
early
early in
in
in
in
in
in
in the
the
the
the
the
the
the visual
visual
visual
visual
visual
visual
visual processing
processing
processing
processing
processing
processing
processing stream.
stream.
stream.
stream.
stream.
stream.
stream.
©
©
©
©
©
©
© 2018
2018
2018
2018
2018
2018
2018 Elsevier
Elsevier
Elsevier
Elsevier
Elsevier
Elsevier
Elsevier Ltd.
Ltd.
Ltd.
Ltd.
Ltd.
Ltd.
Ltd. All
All
All
All
All
All
All rights
rights
rights
rights
rights
rights
rights reserved.
reserved.
reserved.
reserved.
reserved.
reserved.
reserved.
1.
1.
1.
1.
1.
1.
1. Introduction
Introduction
Introduction
Introduction
Introduction
Introduction
Introduction
The
The
The
The
The
The
The number
number
number
number
number
number
number sense
sense
sense
sense
sense
sense
sense describes
describes
describes
describes
describes
describes
describes our
our
our
our
our
our
our ability
ability
ability
ability
ability
ability
ability to
to
to
to
to
to
to estimate
estimate
estimate
estimate
estimate
estimate
estimate the
the
the
the
the
the
the
number
number
number
number
number
number
number of
of
of
of
of
of
of objects
objects
objects
objects
objects
objects
objects or
or
or
or
or
or
or events
events
events
events
events
events
events without
without
without
without
without
without
without counting
counting
counting
counting
counting
counting
counting them
them
them
them
them
them
them (
(
(
(
(
(
(Dehaene,
Dehaene,
Dehaene,
Dehaene,
Dehaene,
Dehaene,
Dehaene,
1997
1997
1997
1997
1997
1997
1997).
).
).
).
).
).
). Symbolic
Symbolic
Symbolic
Symbolic
Symbolic
Symbolic
Symbolic mathematics
mathematics
mathematics
mathematics
mathematics
mathematics
mathematics is
is
is
is
is
is
is unique
unique
unique
unique
unique
unique
unique to
to
to
to
to
to
to educated
educated
educated
educated
educated
educated
educated adult
adult
adult
adult
adult
adult
adult
humans,
humans,
humans,
humans,
humans,
humans,
humans, but
but
but
but
but
but
but we
we
we
we
we
we
we share
share
share
share
share
share
share the
the
the
the
the
the
the number
number
number
number
number
number
number sense
sense
sense
sense
sense
sense
sense with
with
with
with
with
with
with infants
infants
infants
infants
infants
infants
infants (
(
(
(
(
(
(Izard,
Izard,
Izard,
Izard,
Izard,
Izard,
Izard,
Sann,
Sann,
Sann,
Sann,
Sann,
Sann,
Sann, Spelke,
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2004;
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to
to
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toNieder,
Nieder,
Nieder,
Nieder,
Nieder,
Nieder,
Nieder, 2016
2016
2016
2016
2016
2016
2016
be
be
be
be
be
be
be encoded
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in
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(IPS)
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(IPS) and
and
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and
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and
and utilized
utilized
utilized
utilized
utilized
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for
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*
*
*
*
*
*
* Corresponding
Corresponding
Corresponding
Corresponding
Corresponding
Corresponding
Corresponding author
author
author
author
author
author
author.
.
.
.
.
.
. 425
425
425
425
425
425
425 South
South
South
South
South
South
South University
University
University
University
University
University
University Avenue,
Avenue,
Avenue,
Avenue,
Avenue,
Avenue,
Avenue, Room
Room
Room
Room
Room
Room
Room 352,
352,
352,
352,
352,
352,
352, Philadelp
Philadelp
Philadelp
Philadelp
Philadelp
Philadelp
Philadelphia,
hia,
hia,
hia,
hia,
hia,
hia, PA
PA
PA
PA
PA
PA
PA 19104,
19104,
19104,
19104,
19104,
19104,
19104, USA
USA
USA
USA
USA
USA
USA Q2
Q2
Q2
Q2
Q2
Q2
Q2.
.
.
.
.
.
.E-mail
E-mail
E-mail
E-mail
E-mail
E-mail
E-mail address:
address:
address:
address:
address:
address:
address: (N.K.
(N.K.
(N.K.
(N.K.
(N.K.
(N.K.
(N.K. DeWind).
DeWind).
DeWind).
DeWind).
DeWind).
DeWind).
DeWind).ndewind@gma
ndewind@gma
ndewind@gma
ndewind@gma
ndewind@gma
ndewind@gma
ndewind@gmail.com
il.com
il.com
il.com
il.com
il.com
il.com
Available
Available
Available
Available
Available
Available
Available online
online
online
online
online
online
online at
at
at
at
at
at
at www.sciencedirect.com
www.sciencedirect.com
www.sciencedirect.com
www.sciencedirect.com
www.sciencedirect.com
www.sciencedirect.com
www.sciencedirect.com
ScienceDirec
ScienceDirec
ScienceDirec
ScienceDirec
ScienceDirec
ScienceDirec
ScienceDirecScienceDirect
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homepage
page
page
page
page
page
page:
:
:
:
:
:
: www.
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cor tex
cor tex
cor tex
cor tex
cor tex
cor tex
cor tex xxx
xxx
xxx
xxx
xxx
xxx
xxx ( 2018 )
( 2 018 )
( 2 018 )
( 2 018 )
( 2 018 )
( 2 018 )
( 2 018 ) 1
1
1
1
1
1
1ee
e
e
e
e
e14
14
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14
14
14
COR
COR
COR
COR
COR
COR
CORTEX2291_proof
TEX2291_proof
TEX2291_proof
TEX2291_proof
TEX2291_proof
TEX2291_proof
TEX2291_proof 16
16
16
16
16
16
16 April
April
April
April
April
April
April 2018
2018
2018
2018
2018
2018
2018 1/14
1/14
1/14
1/14
1/14
1/14
1/14■
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■
■
■
https://doi.org/10.1016/j.cortex.2018.03.027
https://doi.org/10.1016/j.cortex.2018.03.027
https://doi.org/10.1016/j.cortex.2018.03.027
https://doi.org/10.1016/j.cortex.2018.03.027
https://doi.org/10.1016/j.cortex.2018.03.027
https://doi.org/10.1016/j.cortex.2018.03.027
https://doi.org/10.1016/j.cortex.2018.03.027
0010-9452/
0010-9452/
0010-9452/
0010-9452/
0010-9452/
0010-9452/
0010-9452/ 2018
2018
2018
2018
2018
2018
2018 Elsevier
Elsevier
Elsevier
Elsevier
Elsevier
Elsevier
Elsevier Ltd.
Ltd.
Ltd.
Ltd.
Ltd.
Ltd.
Ltd. All
All
All
All
All
All
All rights
rights
rights
rights
rights
rights
rights reserved.
reserved.
reserved.
reserved.
reserved.
reserved.
reserved.©
©
©
©
©
©
©
Please
Please
Please
Please
Please
Please
Please cite
cite
cite
cite
cite
cite
cite this
this
this
this
this
this
this article
article
article
article
article
article
article in
in
in
in
in
in
in press
press
press
press
press
press
press as:
as:
as:
as:
as:
as:
as: DeWind,
DeWind,
DeWind,
DeWind,
DeWind,
DeWind,
DeWind, N.
N.
N.
N.
N.
N.
N. K.,
K.,
K.,
K.,
K.,
K.,
K., et
et
et
et
et
et
et al.,
al.,
al.,
al.,
al.,
al.,
al., Numerical
Numerical
Numerical
Numerical
Numerical
Numerical
Numerical encoding
encoding
encoding
encoding
encoding
encoding
encoding in
in
in
in
in
in
in early
early
early
early
early
early
early visual
visual
visual
visual
visual
visual
visual cortex,
cortex,
cortex,
cortex,
cortex,
cortex,
cortex, Cortex
Cortex
Cortex
Cortex
Cortex
Cortex
Cortex (2018),
(2018),
(2018),
(2018),
(2018),
(2018),
(2018), https://doi.
https://doi.
https://doi.
https://doi.
https://doi.
https://doi.
https://doi.org/10.1016/
org/10.1016/
org/10.1016/
org/10.1016/
org/10.1016/
org/10.1016/
org/10.1016/j.cortex.2018.03.027
j.cortex.2018.03.027
j.cortex.2018.03.027
j.cortex.2018.03.027
j.cortex.2018.03.027
j.cortex.2018.03.027
j.cortex.2018.03.027
-
decision-making
decision-making
decision-making
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processes
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Miller,
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Miller,
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that
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(Roitman,
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Brannon,
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&
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Platt,
Platt,
Platt,
Platt,
Platt,
Platt,
Platt, 2007
2007
2007
2007
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2007).
).
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). Functional
Functional
Functional
Functional
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Functional imaging
imaging
imaging
imaging
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studies
studies
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Dumoulin,
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visual cortex
cortex
cortex
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arose
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electroencephalography
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Woldorff,
Woldorff,
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Park,
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Park,
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numbers
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et
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al.,
al.,
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al.,
al.,
al., 2013;
2013;
2013;
2013;
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