Directional Visual Motion Is Represented in the Auditory - IA de Investigación especializada en el MIT

Directional Visual Motion Is Represented in the Auditory
and Association Cortices of Early Deaf Individuals

Talia L. Retter1,2, Michael A. Webster1, and Fang Jiang1

Abstracto

■ Individuals who are deaf since early life may show enhanced
performance at some visual tasks, including discrimination of
directional motion. The neural substrates of such behavioral en-
hancements remain difficult to identify in humans, a pesar de
neural plasticity has been shown for early deaf people in the
auditory and association cortices, including the primary audi-
tory cortex (PAC) and STS region, respectivamente. Aquí, we inves-
tigated whether neural responses in auditory and association
cortices of early deaf individuals are reorganized to be sensitive
to directional visual motion. To capture direction-selective re-
sponses, we recorded fMRI responses frequency-tagged to the
0.1-Hz presentation of central directional (100% coherent
random dot) motion persisting for 2 sec contrasted with non-
directional (0% coherent) motion for 8 segundo. We found direction-
selective responses in the STS region in both deaf and hearing

Participantes, but the extent of activation in the right STS region
era 5.5 times larger for deaf participants. Minimal but signifi-
cant direction-selective responses were also found in the PAC
of deaf participants, both at the group level and in five of six
individuals. In response to stimuli presented separately in the
right and left visual fields, the relative activation across the right
and left hemispheres was similar in both the PAC and STS
region of deaf participants. Notablemente, the enhanced right-
hemisphere activation could support the right visual field
advantage reported previously in behavioral studies. Taken to-
juntos, these results show that the reorganized auditory corti-
ces of early deaf individuals are sensitive to directional motion.
Speculatively, these results suggest that auditory and associa-
tion regions can be remapped to support enhanced visual
actuación. ■

INTRODUCCIÓN

The absence of sensory inputs from one modality early in
life has been linked to enhancement of the other senses.
Respectivamente, congenitally deaf people have been shown
to display better performance at some visual tasks than
hearing individuals (p.ej., Shiell, Champoux, & Zatorre,
2014; Bottari, Nava, Ley, & Pavani, 2010; Dye, Hauser,
& Bavelier, 2009; Lore & Song, 1991; Neville & Lawson,
1987; Parasnis & Samar, 1985). Por ejemplo, an enhance-
ment at detecting and discriminating directional visual
motion has been reported in early deaf people (Shiell
et al., 2014; Hauthal, Sandmann, Debener, & Thorne,
2013; in the right visual field [RVF] solo: Bosworth,
Petrich, & Dobkins, 2013; Bosworth & Dobkins, 1999;
Neville & Lawson, 1987). From an ecological perspective,
the daily importance of visual motion may be increased
for deaf individuals, especially for monitoring the periph-
eral visual field, Por ejemplo, when using sign language
(Codina, Pascalis, Baseler, Levin, & Buckley, 2017).
Sin embargo, for other potentially useful visual tasks, No
differences or a decrease in performance has been

This paper is part of a Special Focus deriving from a symposium
en el 2017 International Multisensory Research Forum (IMRF).

1University of Nevada, Reno, 2Universidad de Lovaina

reported across deaf and hearing people (for reviews
on this controversy, see Pavani & Bottari, 2012; mitchell
& Maslin, 2007; Bavelier, Dye, & Hauser, 2006; Parasnis,
1983). The prevalent hypothesis explaining these differ-
ences regards neural plasticity, eso es, the recruitment
of brain areas processing the deprived sense or the reor-
ganization of brain areas processing the existent senses
or engaging in multisensory integration. It is thought that
neural plasticity could support compensatory behavioral
abilities, but only when the underlying functional organi-
zation of the incoming sense is compatible with those
areas (p.ej., Bola et al., 2017; Pascual Leone & hamilton,
2001). Sin embargo, the capacity for neural plasticity of early
deaf individuals to support behavioral advantages in
visual tasks, including those involving motion, has not
been clearly demonstrated.

Extensive neural plasticity has been reported for deaf
individuals’ responses to visual motion. Most strikingly,
several human neuroimaging studies have reported ac-
tivation in the primary auditory cortex (PAC) of deaf
participants in response to moving or flickering visual
estímulos, most often presented in or toward the visual pe-
riphery (peripheral moving dot pattern: Finney, Fine, &
Dobkins, 2001; flickering patch of a full-field luminance
grating: Finney, Clementz, Hickok, & Dobkins, 2003; pe-
ripheral moving dot pattern: Fine, Finney, Boynton, &
Dobkins, 2005; flickering point lights in the RVF: Scott,

Revista de neurociencia cognitiva 31:8, páginas. 1126–1140
doi:10.1162/jocn_a_01378

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Karns, dow, stevens, & Neville, 2014). In addition to au-
ditory cortex, in the multisensory STS region (a term
used to include the STS and adjacent cortex of the supe-
rior and middle temporal gyrus and angular gyrus;
alison, Chip, & McCarthy, 2000), a trend has been shown
for higher activation and significantly more pronounced
attentional enhancement in deaf people in response to
visual dot motion (Bavelier et al., 2001). In the study by
Scott et al. (2014), a larger area of activation around the
STS was reported in deaf participants, including the pos-
terior superior and middle temporal gyrus. Changes in
responsiveness to peripherally presented visual motion
or flickering stimuli have also been reported in human
visual area hMT+: Increased (left-hemisphere) activación
and/or extent of activation has been reported in deaf
gente (Scott et al., 2014; Bavelier et al., 2000, 2001;
but see also Fine et al., 2005). To a lesser degree, otro
areas implying cross-modal neuroplasticity for motion or
flicker in the early deaf people include the posterior pa-
rietal cortex, anterior cingulate, and FEF/supplementary
eye field (Scott et al., 2014; Bavelier et al., 2001).

De nuevo, sin embargo, the relationship between such neural
plasticity in early deaf people and behavioral advantages
in visual motion detection or discrimination has not been
well documented. Recent evidence from animal studies
suggests a causal link between reversible lesions in the
auditory cortex and behavioral advantages at visual lo-
calization and movement detection in cats (Lumbar,
Meredith, & Kral, 2010; see also Meredith et al., 2011).
Yet for humans, only noninvasive, correlative evidence
has been provided. Structurally, Por ejemplo, correlations
have been found for deaf individuals between the relative
amount of auditory cortex (planum temporale) or visual
corteza (V1) devoted to processing peripheral motion and
behavioral performance in motion detection tasks (audi-
conservador: Shiell, Champoux, & Zatorre, 2016; visual: Levin,
Codina, Buckley, de Sousa, & Baseler, 2015). Suggestive
evidence has also been provided by showing that the re-
cruitment of reorganized brain regions in early deaf indi-
viduals shows selective responses to a visual task for
which there is behavioral enhancement. Por ejemplo,
four cardinal locations of visual stimuli could be decoded
from the auditory cortex in deaf individuals with neuro-
imaging, suggesting that representations in the auditory
cortex align with those in the visual cortex (Almeida et al.,
2015). Aquí, we aim to add to these findings by asking
whether deaf individuals’ enhanced ability in speed
and/or accuracy at discriminating the direction of visual
motion could be supported by direction-selective re-
sponses in brain areas evidencing neural plasticity.

Directional visual motion is a particularly salient visual
stimulus and is known to selectively activate a subset of
areas in the neurotypical human brain responding to vi-
sual motion more generally. Strong direction-selective
responses have been found in human visual area hMT
+/ V5 (p.ej., Huk, Ress, & Heeger, 2001; Morrone et al.,
2000; Tootell et al., 1995). Other implicated areas include

V3/V3A and, en un grado menor, the rest of V1–V4 (cervezas &
Norcia, 2009; Huk et al., 2001; Tootell et al., 1995; finding
large effects also in V1 with EEG source imaging). El
representation of directional motion within these cortical
areas was first revealed by single-cell recordings in mon-
keys, reporting columnar direction tuning (Felleman &
VanEssen, 1987; Albright, 1984; Dubner & Zeki, 1971;
Hubel & Wiesel, 1961). Desafortunadamente, debido a la
spatial scale, such direction tuning cannot be studied
noninvasively in humans, and direction-specific represen-
tation in humans has thus remained elusive (see Kamitani
& Tong, 2006, for a potential exception, but also Beckett,
Peirce, Sanchez-Panchuelo, Francisco, & Schluppeck, 2012;
for axis of motion mapping at 7 t, see Zimmermann
et al., 2011). Despite this, direction-selective areas may
be identified in the human brain with fMRI with stim-
ulation presentation techniques, such as contrasting
directional (es decir., coherent) motion with directionless (es decir.,
noncoherent) motion or dynamic noise (Morrone et al.,
2000; Beauchamp, Cox, & DeYoe, 1997; Braddick,
Hartley, Atkinson, Wattam-Bell, & Tornero, 1997; in EEG/
magnetoencephalography: Palomares, cervezas, Wade,
Cottereau, & Norcia, 2012; cervezas & Norcia, 2009; Nakamura
et al., 2003; Lam et al., 2000; tyler & Kaitz, 1977).

Aquí, we used a sensitive approach to investigate the
spatial extent and activation of direction-selective brain
regions in early deaf and hearing people, focusing on
the auditory and association cortices, the PAC and STS
región, respectivamente, in comparison with visual area hMT+.
Específicamente, we used fMRI together with a frequency-
tagging approach (p.ej., gao, Gentile, & rossión, 2017;
Koening-Robert, VanRullen, & Tsuchiya, 2015; Ernst,
Boynton, & Jazayeri, 2013; Morrone et al., 2000; ángel,
zhang, & Wandell, 1997; Chip, alison, Sangre, & McCarthy,
1995; Bandettini, Jesmanowicz, Wong, & Hyde, 1993) a
identify periodic changes from noncoherent to direc-
tional random-dot motion (Morrone et al., 2000; see also
Palomares et al., 2012; cervezas & Norcia, 2009; Atkinson
et al., 2008). We were thus able to acquire signals with
a high signal-to-noise ratio that were independent of a
hemodynamic response function model. By using a
contrast of directionless-to-directional motion, we were
also able to capture direction-selective responses (nota
that these responses are not direction specific) locked
precisely to the frequency of coherence onset. To follow
up on a behavioral advantage for direction discrimina-
tion typically reported in the RVF for deaf individuals
(Bosworth et al., 2013; Bosworth & Dobkins, 1999;
Neville & Lawson, 1987), we presented visual stimuli
in the left visual field (LVF) and RVF as well as centrally.
When activation was found, we further explored potential
qualitative differences across hearing and deaf individuals
in terms of spatial extent, RVF versus LVF response, y
hemispheric lateralization. Juntos, these comparisons
allowed us to assess the potential neural bases of en-
hanced visual motion processing reported in previous
studies for early deaf people.

Platos, Webster, and Jiang

1127

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MÉTODOS

Participantes

Two groups of participants, early deaf and hearing controls,
were tested in the experiment, which was approved by the
institution review board of the University of Nevada, Reno,
and conducted in accordance with the Code of Ethics of
the World Medical Association (Declaration of Helsinki).
Each group consisted of six adults, recruited from northern
Nevada and California. Our deaf participants included those
who experienced severe-to-profound sensorineural hearing
loss at an early age. They had no ability to understand
auditory speech but were proficient in sign language (ver
Mesa 1 for deaf participants’ details). The mean age of deaf
participants was 36 años (DE = 8.2, range = 26–49 years);
the mean age of hearing participants was 33 años (DE =
8.5, range = 26–48 years). Four of the hearing and one
of the deaf participants were male; one hearing participant
was left-handed. All participants were unaware of the ex-
perimental design, except for one hearing participant,
who was author T. l. R. All participants reported visual
acuity in the normal or corrected-to-normal range.

f MRI Acquisition

f MRI scanning was performed with a 3-T Philips Ingenia
scanner using a 32-channel digital SENSE head coil (Philips
Medical Systems) at the Renown Regional Medical Center,
Reno, NV. Volumetric anatomical images were acquired at a
resolution of 1 mm3 using a T1-weighted magnetization
prepared rapid gradient echo sequence. Functional BOLD
signals were acquired through a continuous design at a
resolution of 2.75 × 2.75 × 3 mm voxels, with no gap. A
repetition time of 2 sec was used to acquire 30 transverse
slices in an ascending order, with an echo time of 17 mseg,
a flip angle of 76°, y un 220 × 220 mm2 field of view.

Visual Motion Stimuli

Visual motion was displayed with random-dot kinemato-
gramos, based on the incremental displacement across

monitor refresh frames of individual dots within a circular
campo (Braddick, 1974; Julesz, 1971; Anstis, 1970). Frames of
white dots against a black background were generated with
a custom script running over MATLAB (The MathWorks),
refreshing at a rate of about 60 Hz, with a 500-msec
lifespan to discourage participants tracking the movement
of individual dots. Given some inconsistency in presenta-
tion timing because of online drawing of dot positions,
the motion display was adapted for precise periodic stimu-
lus presentation by exporting generated dot motion frames
and then displaying them at a precisely controlled periodic
tasa de 60 Hz using custom software running over Java.
Viewed on the testing monitor, the stimulus field diame-
ter subtended 8.5° of visual angle, with individual dots
subtending 1.35 en. in diameter, moving at a speed of
3.4°/sec, at a density of 12.5 dots/deg.

We created directional stimulus sequences in four direc-
ciones (arriba, bien, abajo, and left) as well as nondirectional,
noncoherent dot motion. To create the directional se-
quences, a 30-sec sequence of 1,800 sequential stimulus
frames creating the appearance of 100% coherent right-
ward visual motion was extracted. The rightward stimuli
frames were rotated by increments of 90° to create se-
quences of downward, leftward, and upward motions, re-
spectively, with minimal variation across directions. En
the functional scans, these sequences each repeated four
times in immediate succession, leading to a block of 2 segundo
of directional motion. To create sequences of non-
directional motion, 100% noncoherent motion stimulus
frames were similarly extracted from 30-sec sequences; a
fill the longer proportion of nondirectional-to-directional
motion duration in the testing sequences with consistent
stimulus update intervals, this procedure was repeated
three additional times. Note that these 30-sec sequence
pieces also served as “incoherent jumps” to prevent a
specific confound of full dot replacement at the onset
and offset times of directional and nondirectional motion
(see the following section; Braddick, Birtles, Wattam-Bell,
& Atkinson, 2005; Wattam-Bell, 1991). In functional scans,
these four nondirectional sequence sets were each re-
peated four times in immediate succession, defining a

Mesa 1. Demographic Information for the Early Deaf Participants

Age ( Años)

Sex

Handedness

Deafness
Acquisition

Cause of Deafness

Auditory Deprivation,
Left/Right (dB)

Signing
Acquisition

METRO

F = female; M = male; R = right.

12 meses

Unknown

15 meses

Fever

Birth

Maternal gestational measles

Genetic (coex26)

Hereditary

Unknown

95/95

Total/85

100/90

85/85

80/70

98/96

12 años

15 meses

11 años

< 1 year < 1 year 1 year 1128 Journal of Cognitive Neuroscience Volume 31, Number 8 l D o w n o a d e d f r o m h t t p : / / d i r e c t . m i t . e d u / j / o c n a r t i c e - p d l f / / / 3 1 8 1 1 2 6 1 7 8 8 6 7 7 / / j o c n _ a _ 0 1 3 7 8 p d . f b y g u e s t t o n 0 7 S e p e m b e r 2 0 2 3 block of 8 sec of nondirectional motion. Participants viewed the stimulation monitor with a mirror attached to the MR head coil. Periodic Visual Stimulation Procedure Functional scans consisted of periodic alternation between directional and nondirectional motion over a duration of 5.1 min. Scans began with 2 sec of a white fixation cross on a black background, followed by a 2-sec fade-in period, in which stimulus luminance contrast gradually increased to 100%. Stimuli were then shown over a duration of 300 sec in a fixed pattern of 2 sec of directional motion followed by 8 sec of nondirectional motion. Periods of directional mo- tion thus onset every 10 sec, leading to a direction-selective frequency-tagged rate of 1/10 sec, that is, 0.1 Hz (Figure 1A). Within each scan, the direction of motion also consistently alternated at each presentation cycle, for example, from upward to downward motion, leading to a direction-specific frequency-tagged rate of 0.05 Hz. Finally, the scans ended with 2 sec of stimulus fade-out and 2 sec of the white fixation cross. Four participants from each of the deaf and hearing groups saw contrasts of up/down and left/right motions (Trial Lists 1 and 2), and the remaining two participants of each group saw contrasts of up/left and right/down motions (Trial Lists 3 and 4). Because no clusters of significant responses to direction-specific motion at 0.05 Hz were found for any participants in any trial lists, data were combined across trial lists within each group to examine the direction-selective response at 0.1 Hz. Visual Field Conditions Scans designed to localize brain regions responding to visual motion contained stimuli presented in a central visual field (CVF) condition. In two additional scan con- ditions designed to measure the amplitude of brain acti- vation, stimuli were presented in either the right or left peripheral visual field. In the CVF condition, stimuli were presented in the center of the stimulation monitor to- gether with a superimposed central fixation cross. From a viewing distance of 134 cm, the monitor supported a field of view of 29° × 17°. Thus, when presented in the right or left peripheral visual field conditions, the stimulus was translated laterally to the edge of the monitor and the fixation cross shifted laterally 4° from center in the opposite direction, so that the distance between the proximal edge of the stimulus and the fixation cross subtended 10° (e.g., Jiang, Beauchamp, & Fine, 2015). Four scan repetitions of each condition were presented sequentially to discourage participants from moving their heads as stimulus location changed. Each participant was presented with every con- dition, leading to 12 scans for a total testing time of about 1 hr. In odd trial lists, scans began with stimuli presented in the CVF, whereas in even trial lists, stimuli were first presented in one of the peripheral visual fields. The order of conditions was identical across participant groups. Behavioral Task Participants were instructed to fixate on the centrally pre- sented white fixation cross. The cross changed shape to a circle for a duration of 200 msec at random intervals (a minimum of 800 msec in between changes) 30 times within each scan, that is, once about every 10 sec. Par- ticipants were asked to use a response box to report the direction of motion at the time of the fixation shape change. This task was designed to facilitate participants’ fixation as well as to encourage attention to the direction of motion of the stimulus. Figure 1. (A) Stimulation sequences consisted of 2 sec of directional (100% coherent dot) visual motion followed immediately by 8 sec of nondirectional (0% coherent dot) motion. The onset of directional motion thus occurred periodically every 10 sec, predicting a direction-selective response in the frequency domain at 0.1 Hz (i.e., 1/10 sec). The arrows drawn on the figure are purely for illustrating the direction of dot motion. (B) Top: An example of the BOLD response recorded by fMRI from a single voxel in visual area hMT+ from a hearing participant, averaged over four runs of visual motion presented in the CVF and DC corrected. Its location is illustrated on the sagittal slice of this participant’s anatomy in Talairach space. Bottom: A fast Fourier transform (FFT) is applied to each voxel to transform the data into the temporal frequency domain. This example voxel is sensitive to directional motion, as evidenced by the high-amplitude BOLD signal of the 0.1-Hz response peak. Retter, Webster, and Jiang 1129 l D o w n o a d e d f r o m h t t p : / / d i r e c t . m i t . e d u / j / o c n a r t i c e - p d l f / / / 3 1 8 1 1 2 6 1 7 8 8 6 7 7 / / j o c n _ a _ 0 1 3 7 8 p d . f b y g u e s t t o n 0 7 S e p e m b e r 2 0 2 3 f MRI Data Analysis Preprocessing. Anatomical and functional data were analyzed with BrainVoyager v20.0 and the BVQXTools toolbox (Brain Innovation B.V.) together with MATLAB R2013b. Functional scan data were imported into BrainVoyager and preprocessed with corrections for slice scan time and 3-D motion (aligned to the first functional scan for intersession alignment). They were temporally filtered with a linear de-trending; no spatial smoothing was applied. Anatomical scans, similarly imported into BrainVoyager, were subjected to an isotropic voxel trans- formation and aligned according to standard anterior and posterior commissure points. For display across par- ticipants, data were transformed into a conventional Talairach space (Talairach & Tournoux, 1988). Func- tional scans were coregistered to each participant’s cor- responding anatomical images. Initial alignment was fine-tuned through an affine transformation and mini- mally corrected with visual inspection. Spatial normaliza- tion of the functional data was applied through a volume time course transformation. Frequency domain processing. The volume time course files of each functional scan were imported into MATLAB for frequency domain analyses. They were cropped to 150 volumes of 2 sec, containing exactly 30 presentation cycles of 0.1-Hz directional motion and ex- cluding the first and last two volumes corresponding to fixation cross and fade-in/out presentation. BOLD data from each participant from the four scans per condition were averaged in time to reduce noise from non-phase- locked activation, that is, from activation not driven by periodic stimulus presentation. A DC correction was ap- plied to remove the mean signal offset, and the data were transformed into a normalized amplitude spectrum through a fast Fourier transform (Figure 1B). The resulting BOLD amplitude spectrum contained a range of 0–0.25 Hz with a frequency resolution of 0.0033 Hz. For each fre- quency bin, x, a baseline range was defined as 20 sur- rounding frequency bins, encompassing a range of about 0.07 Hz centered around x. To assess significance during CVF scans of the 0.1-Hz response at each voxel, z scores were generated by subtracting from x the mean baseline value and dividing the result by the standard deviation of the baseline. To display BOLD response amplitudes in pre- determined regions (see section below) during RVF and LVF scans, baseline-subtracted amplitude values were similarly generated by subtracting the mean baseline value from x (e.g., Retter & Rossion, 2016). The resulting files were reimported into BrainVoyager for display. ROIs. Given previously reported findings of neural plas- ticity in the PAC and association auditory cortex in deaf individuals (e.g., Scott et al., 2014; Karns, Dow, & Neville, 2012; Fine et al., 2005; Finney et al., 2001, 2003) and direction-selective responses in visual area hMT+ (e.g., Huk et al., 2001; Morrone et al., 2000; Tootell et al., 1995), we a priori focused our analyses on the PAC and STS region, potentially including part of the STS/PT, middle temporal gyrus, and angular gyrus (Allison et al., 2000; see also Scott et al., 2014, for activation in deaf participants), and hMT+. To define the STS region and hMT+, we used a func- tional cluster-based criterion from direction-selective responses at 0.1 Hz to motion presented in the CVF (clusters > 150 vóxeles). Significance thresholding was ap-
plied at the individual participant level (range: z > 2.6 a
z > 5.7), to approximately equalize the number of signif-
icant voxels across commonly active regions, incluido
hMT+ (six deaf and six hearing, in at least one hemi-
sphere), the STS region (six deaf and six hearing), early
visual areas (six deaf and six hearing), and the lateral oc-
cipital complex (five deaf and six hearing). In relevant
casos, the threshold was increased for two regions, ap-
plied bilaterally, to spatially separate them. The mean
total voxel number across participants after thresholding
era 15,138 voxels and did not differ significantly across
grupos (deaf: m = 13,636, SE = 1,422; hearing: m =
16,641, SE = 1,583), t= 1.41, pag = .19, re = 0.73 (two-
cola).

In a separate analysis, we defined the PAC, a region
that cannot be functionally defined in deaf participants,
using the Julich probabilistic atlas in the SPM Anatomy
Toolbox (Eickhoff, Heim, Zeilles, & Amunts, 2006;
Eickhoff et al., 2005). Following a procedure described
in Eickhoff et al. (2006), we included the volume assign-
ment to all subregions for PAC (Morosan et al., 2001) en
the summary map of all areas (maximum probability
map). This procedure ensured no overlap between any
two cytoarchitectonic defined areas. The PAC ROI was
then transformed to Talairach space and applied to each
participant’s brain volume. It was further separated into
left and right hemisphere PAC for each participant.

Statistical tests. For the functionally defined ROIs,
a saber, the STS region and hMT+, we investigated
whether there were significant differences in the spatial
extent and amplitude of activation between the deaf and
hearing participant groups. The spatial extent and ampli-
tude of the STS region and hMT+ were thus compared
across the deaf and hearing participant groups with non-
parametric Mann–Whitney U tests, given the relatively
small sample size. To compare differences in the spatial
extent of the STS region and hMT+, the number of sig-
nificant voxels was used. To compare the amplitude dif-
ferences in these ROIs to stimuli presented in the LVF
and RVF, baseline-subtracted amplitude values at 0.1 Hz
for each participant were averaged across voxels within
their individually defined ROIs for the LVF and RVF re-
sponses separately. When a cluster-based ROI could not
have been defined in one hemisphere (STS: two deaf and
one hearing in the left hemisphere; MONTE: one deaf partic-
ipant in the left hemisphere), no corresponding ampli-
tude values were included in the analysis.

1130

Revista de neurociencia cognitiva

Volumen 31, Número 8

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For the probabilistically defined ROI, eso es, the PAC,
we investigated whether there were significant responses
in the deaf and/or hearing participants. To determine
response significance in the PAC ROI, an amplitude spec-
trum was computed from the averaged BOLD responses
to motion presented in the CVF of all bilateral PAC voxels
across participants in each group. z Scores were then
calculated on this averaged spectrum with a threshold
of p < .001 (z > 3.10) for significance for this sensitive
group-level analysis. Given some debate about whether
PAC responses occur only as a result of group level aver-
aging (p.ej., as shown in Finney et al., 2001; but see Scott
et al., 2014), significance was also assessed similarly at
the individual participant level, with the typical thresh-
old of p < .05 (z > 1.64). To compare the number of
significantly direction-selective voxels across the PAC
and the STS region, the PAC ROI was thresholded at
the individual level defined previously for demarcating
the STS region (es decir., encompassing a range of z > 2.6
to z > 5.7).

RESULTADOS

The STS Region and Visual Middle
Temporal Complex

The centrally presented visual motion trials were used to
localize direction-selective responses in deaf and hearing
individuals. These responses were frequency-tagged at
0.1 Hz, eso es, the rate at which directional (100% dot
coherencia) motion onset (and continued for 2 segundo) im-
mediately after 8 sec of directionless motion (0% dot
coherencia).

m = 3,591 mm3, SE = 596.2; hearing: m = 653 mm3,
SE = 22.6), with no pronounced differences in the left
hemisferio (deaf: m = 294 mm3, SE = 164.8; hearing:
m = 315 mm3, SE = 124.8; Figura 2A). Statistically, este
led to a significant difference in the extent of STS
region activation across participant groups in the right
STS region only: U = 1, pag = .004 (left STS: U = 36, pag =
.70).

The STS region in the right hemisphere was centered
at Talairach x = 54, y = −42, and z = 9 for deaf partic-
ipants and x = 55, y = −39, and z = 16 for hearing par-
ticipants (for individual regions, ver figura 3). El
location of the STS region was particularly reliable in
the right hemisphere for deaf participants; The range of
its center Talairach x coordinates (ver figura 3) was x =
52–58 (SE = 0.84) for deaf participants, comparado con
x = 48–66 (SE = 3.07) for hearing participants (en el
left hemisphere, the range was x = 45–61 [SE = 3.45]
for deaf participants and x = 50–65 [SE = 2.96] for hear-
ing participants).

The area of hMT+ did not appear to differ greatly across
participant groups, although the right hemisphere (deaf:
m = 1,847 mm3, SE = 183.3; hearing: m = 1,304 mm3,
SE = 321.2) appeared larger than the left (deaf: m =
1,033 mm3, SE = 365.9; hearing: m = 1,335 mm3, SE =
291.1) for deaf participants only (Figura 2B). Sin embargo,
statistically, there was not a significant difference in the
extent of hMT+ activation across deaf and hearing partic-
ipants, in either the right, U = 11, pag = .31, or left, U = 14,
pag = .59, hemisferio. En resumen, the only significant dif-
ference found between deaf and hearing participants in
terms of area of activation was a greater extent of the right
STS region for deaf participants.

Direction-selective Responses Are More Extensive in the
Right STS Region for Deaf Participants

Responses in the LVF vs. RVF: An RVF STS Region Bias for
Deaf Participants

The area of the STS region was 5.5 times larger in deaf
than hearing individuals in the right hemisphere (deaf:

The amplitude of responses within the STS region and
hMT+ ROIs identified previously was used to quantify

Cifra 2. The size of STS region and hMT+ ROIs in deaf and hearing individuals. In the center, a sagittal slice (hemisferio derecho; Talairach x = 46)
provides an example of the location of these regions defined in a single hearing participant; the STS region is drawn in green, being dorsal and slightly
more anterior relative to hMT+, drawn in blue/purple. (A) The extent of activation in the STS region in deaf and hearing individuals (promedio [Avg.]
across groups plotted on the far right, with error bars representing ± 1 SE ), in each of the left and right hemisphere. (B) The extent of activation in
hMT+, plotted as in A.

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Cifra 3. The STS region ROIs in the anatomy of deaf and hearing individuals, in Talairach space. Data are thresholded with individually
defined z score values (see the scale in the top right corner). These sagittal slices are centered around the functionally defined ROI for each
hemisferio; in three cases where a functional ROI could not be defined in one hemisphere, the x coordinate mirrors that of the other
hemisferio (and is presented in italics).

the response to visual motion presented in separate trials
in the LVF and RVF. We expected that the larger size of
the right STS region only in deaf individuals might be
accompanied by enhanced activity in response to visual
stimuli presented in the LVF. En cambio, the results showed
the opposite, eso es, that STS region responses of deaf
individuals were of larger amplitude for stimuli presented
in the RVF than LVF (Figura 4A). En efecto, for stimuli
presented in the RVF, there was a significantly higher
response for deaf than hearing participants in the right
STS, U = 0, pag = .002, which only neared significance in
the left STS, U = 2, pag = .063. A diferencia de, for stimuli pre-
sented in the LVF, there were no significant differences
across participant groups—right STS: U = 9, pag = .18; izquierda
STS: U = 9, pag = .91.

In hMT+, the pattern of amplitude responses to stim-
uli in the LVF and RVF appeared highly similar across
the left and right hemispheres for deaf and hearing

Participantes (Figura 4B). This pattern was described by
a contralateral visual field to hemisphere advantage, par-
ticularly for the RVF/left hemisphere (see Figure 4B).
Across participant groups, sin embargo, there were no sig-
nificant differences for stimuli presented in either the
RVF (right hMT+: U = 12, pag = .39; left hMT+: U =
14, pag = .93) or LVF (right hMT+: U = 15, pag = .70; izquierda
hMT+: U = 15, pag = 1.0). En general, there were thus no
differences between deaf and hearing participants in
hMT+ responses to directional motion in the LVF or
RVF, but deaf participants had more activation than
hearing participants in the right STS region to stimuli
presented in the RVF.

The PAC

The PAC was defined with a probabilistic atlas for both
deaf and hearing participants. To determine response

Cifra 4. Amplitude values (baseline-subtracted) of the frequency domain analysis of periodic BOLD signal changes to directional motion at
0.1 Hz. Data are reported for deaf and hearing participants in response to visual stimuli presented in the LVF and RVF in the left (LH) y correcto (RH)
hemispheres. Results are shown in the ROIs defined previously for the STS region (A) and hMT+ (B). Group data are plotted in bar graphs
(error bars plotting ± 1 SE ), and individual data are plotted as superimposed dots (deaf, filled-in; hearing, unfilled-in); each participant is plotted in a
consistent color across plots.

1132

Revista de neurociencia cognitiva

Volumen 31, Número 8

significance in this region, an amplitude spectrum was
derived from the averaged BOLD responses to motion
presented in the CVF of all PAC voxels across participants
for each group.

Significant But Minimal Direction-selective Responses in
the PAC for Deaf Participants

At the individual participant level, direction selectivity at
0.1 Hz was evidenced in five of six deaf participants in the
bilateral PAC (zs ranging from 1.79 a 3.19, ps < .05). A significant direction-selective response emerged at 0.1 Hz for the deaf participants across the bilateral audi- tory cortex (z = 5.80, p < .0001; right PAC: z = 7.78; left PAC: z = 3.40; Figure 5A). A direction-selective response was not found in five of six hearing participants (all zs < 1.34, ps > .05); sin embargo,
a significant response was found in one hearing par-
ticipant who was not naive to the experimental design
(z = 4.36, pag < .0001). When including all six participants of the hearing group, the direction-selective response in the bilateral PAC reached significance at a threshold of p < .05 (bilaterally: z = 2.32, p = .010; right PAC: z = 1.88; left PAC: z = 2.27); when removing the nonnaive participant, the PAC was not significant in the hearing group (bilaterally: z = 1.39, p = .082). Despite significant responses in the PAC in deaf partic- ipants, the extent of direction-selective responses was minimal, with significant voxels subtending only 14.1% of the bilateral PAC area (at z > 3.10, pag < .001) for deaf participants at the group level (i.e., grand-averaged am- plitude spectra; see Methods). When the lowest and highest z score thresholds applied for individual partici- pants (z > 2.6 to z > 5.7) were applied to the group level
datos, the percentage of significant bilateral PAC area at
the group level ranged from, respectivamente, 21.8% a
0.67%. To put this in perspective, this area was more than
28 times smaller than the extent of activation in the bilat-
eral STS region for deaf participants when defined at the
same significance threshold for individual participants
(see Figure 5B; mean PAC area: 1.11%, SE = 0.43%). En
summary, significant PAC responses were present but
minimal in the deaf participant group.

Responses in the LVF vs. RVF: The PAC Hemispheric
Activation Mirrors the STS

The responsivity of the BOLD amplitude of direction-
selective responses to visual motion presented in the
RVF or LVF was investigated, as in the Responses in the
LVF vs. RVF: An RVF STS Region Bias for Deaf Participants
sección (Figure 5C). The resultant pattern of activation
across hemispheres in the PAC was reminiscent of that
of the STS region (compare Figure 5C with Figure 4A).
Note that the large amplitude differences across the
STS region and PAC are not comparable directly, porque
of the different methods of definition of these regions
(es decir., the STS region was defined functionally to include
only significant voxels, whereas the PAC was defined as
all voxels within a predefined region).

Cifra 5. Responses to directional visual motion activated the PAC of deaf individuals. (A) A nivel de grupo, areas of activation in deaf participants’
temporal lobes encompassed the probabilistic area of the auditory cortex (shown in light blue on the standard Colin 27 cerebro; datos
at p < .001). Moreover, (B) significant responses to direction-selective motion at 0.1 Hz were found in the PAC (shaded in light blue) at the individual level, although the area of activation was small relative to the STS region: z Scores of three deaf individuals are shown here at the same thresholded level used to define their individual STS ROI (D2: z > 4.57; D3: z > 5.7; D4: z > 2.6). (C) The pattern of activation in the left and
right PAC for deaf participants to visual motion in the LVF and RVF was similar to that of their STS region (compare with Figure 4A). De nuevo, group data
are plotted in bar graphs (error bars plotting ± 1 SE ), and individual data are plotted as superimposed dots; colors are consistent across plots
and labeling in B. L = left; R = right.

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DISCUSIÓN

We used an fMRI frequency-tagging approach to identify
direction-selective brain regions in early deaf and hearing
gente, investigating the spatial extent of their activation
(in response to stimuli presented in the CVF) y el
amplitude of their activation (in response to stimuli
presented separately in the LVF and RVF). We focused
our analysis on the PAC and associative STS region, en
comparison with visual area hMT+. We predicted that
direction-selective response would be found in the PAC
and STS region, in line with enhanced behavioral abili-
ties reported for early deaf individuals in discriminating
and/or detecting directional visual motion (Shiell et al.,
2014; Bosworth et al., 2013; Hauthal et al., 2013; Bosworth
& Dobkins, 1999; Neville & Lawson, 1987). Tenga en cuenta que
we are able to identify direction-selective responses
emerging from a contrast of directional versus nondirec-
tional visual motion in our frequency-tagging paradigm.
Direction-selective motion responses are more selective
than motion-selective responses but less selective than
direction-specific (p.ej., leftward-selective) respuestas. On
the other hand, previous studies investigating motion-
related responses in the early deaf people have reported
motion-selective, rather than direction-selective, re-
sponses (p.ej., Fine et al., 2005; Finney et al., 2001).

Direction-selective Responses Are Found in the STS
Region for Both Hearing and Deaf Individuals

To our knowledge, this is the first study showing di-
rection selectivity for translational visual motion in the
human STS region (see Figures 2A and 3), here encom-
passing the posterior to middle STS, superior temporal
gyrus, and middle temporal gyrus (for direction selectiv-
ity with rotational head and ellipsoid motion, see Carlin,
Rowe, Kriegeskorte, Thompson, & Calder, 2012). El
STS region is known to respond to visual (biological)
motion in neurotypical humans and nonhuman animal
modelos (para una revisión, see Allison et al., 2000; see also,
p.ej., Noguchi, Kaneoke, Kakigi, Tanabe, & Sadato, 2005;
Grossman & Blake, 2001). Además, direction-selective
tuning of single neurons to visual motion has been re-
ported in the STS region of monkeys (p.ej., Nelissen,
Vanduffel, & Orban, 2006; Oram, Perrett, & Hietanen,
1993; bruce, Desimone, & Bruto, 1981; Zeki, 1978).
The absence of direction-selective STS responses in past
human neuroimaging or source localization studies may
be for several reasons: Por ejemplo, these studies fo-
cused on more traditionally, retinotopically defined
areas, and there may be differences in activation resulting
from the directional/nondirectional motion contrast used
here and the motion adaptation paradigms favored previ-
iosamente. Tenga en cuenta que, in previous studies, direction-selective
responses were reported only in visual areas V1 through
hMT+/ V5 and the lateral occipital complex (hong, Tong,
& Seiffert, 2013; cervezas & Norcia, 2009; Huk et al., 2001;

Tootell et al., 1995). Además, the frequency-tagging
paradigm applied here may have provided methodological
advantages, enabling a powerful contrast of directional and
nondirectional motion, an analysis with a high signal-to-
noise ratio, and not relying on a hemodynamic response
function model (p.ej., Gao et al., 2017; Koening-Robert
et al., 2015; Ernst et al., 2013; Morrone et al., 2000;
Engel et al., 1997; Puce et al., 1995; Bandettini et al., 1993).
The direction-selective STS region could be function-
ally defined in all individual deaf and hearing participants
in the right hemisphere and in five deaf and four hearing
participants in the left hemisphere (ver figura 3). Fue
2–12 times larger in the right than left hemisphere, para
the hearing and deaf participants, respectivamente (ver
Figura 2A). A nivel de grupo, in the right hemisphere,
this region was centered at Talairach coordinates of x =
55, y = −39, and z = 16 for hearing participants and x =
54, y = −42, and z = 9 for deaf participants. The local-
ization of the STS region here is similar to that reported
in previous studies (p.ej., for deaf participants, respuesta
to visual motion: x = 56, y = −40, z = 8, en mesa 5 de
Bavelier et al., 2001; for neurotypical participants in
response to visual, tactile, and auditory stimuli: left ante-
rior inferior coordinates of x = 52, y = 44, z = 15, en
Beauchamp, Yasar, Frye, & Ro, 2008).

This STS region also showed a right-hemisphere ad-
vantage in terms of response amplitude to stimuli shown
in the LVF and RVF, particularly for deaf participants. En
contrast, there was no left-hemisphere advantage appar-
ent for stimuli shown in the RVF for either participant
grupo (see Figure 4A). These results are in line with
larger responses to visual motion in the right hemisphere
generally (p.ej., Corballis, 2003; Finney et al., 2001;
Kubova, Kuba, Hubacek, & Vit, 1990; see also Weeks
et al., 2000, for an example of right-hemisphere domi-
nance to auditory motion in congenitally blind partici-
pants) as well as interhemispheric transfer of visual
motion information (Brandt, Esteban, Bense, Yousry, &
Dieterich, 2000; see also Motter, Steinmetz, Duffy, &
Mountcastle, 1987) and previous reports of no contralat-
eral organization in the STS region (p.ej., Grossman &
Blake, 2001; see also Saygin & Sereno, 2008).

The PAC Shows Direction-selective Visual Motion
Responses in Early Deaf Individuals

We discovered significant direction-selective responses
to visual motion in a probabilistically defined PAC region
in early deaf people (ver figura 5). The extent of this ac-
tivation was highly dependent on the significance thresh-
old used; at p < .001, it appeared to cover 14.1% of the bilateral PAC for the early deaf group. In comparison with the extent of activation in the STS at the same signifi- cance threshold, this area is more than 28 times smaller. Nevertheless, when averaging across all voxels in the bi- lateral PAC, a significant response emerged for five of six deaf participants ( p < .05). 1134 Journal of Cognitive Neuroscience Volume 31, Number 8 l D o w n o a d e d f r o m h t t p : / / d i r e c t . m i t . e d u / j / o c n a r t i c e - p d l f / / / 3 1 8 1 1 2 6 1 7 8 8 6 7 7 / / j o c n _ a _ 0 1 3 7 8 p d . f b y g u e s t t o n 0 7 S e p e m b e r 2 0 2 3 Responses to visual stimuli were first reported in the PAC for early deaf people in response to peripheral mov- ing dots at the group level (Finney et al., 2001). PAC ac- tivation was replicated in the early deaf people in response to moving or flickering stimuli, most often in or near the visual periphery (Finney et al., 2001, 2003). Importantly, these results were likely not an effect of group averaging or imprecise PAC definition: A recent study identified PAC activation defined anatomically at the individual participant level, using the transverse tem- poral gyrus, also known as Heschl’s gyrus, with flickering point lights in the RVF (Scott et al., 2014). In this study, the amount of activation in the PAC was reported only in comparison for peripherally versus perifoveally pre- sented flicker dots, preventing a direct comparison with the extent or amount of activation reported here. Still, our finding that PAC activation is—at least to some extent—direction selective adds to our knowledge of neural plasticity in this region for early deaf people. The pattern of PAC activation in response to stimuli presented in the RVF and LVF is highly reminiscent of that of the STS region (see Figures 5C and 4A). One pos- sibility is that the PAC projects information into the STS region, a sensory association area (e.g., Hackett et al., 2007; Smiley et al., 2007; Seltzer et al., 1996; Seltzer & Pandya, 1978; Benevento, Fallon, Davis, & Rezak, 1977; see also Beauchamp et al., 2008). In addition, the STS also projects information back to the superior temporal gyrus (e.g., Barnes & Pandya, 1992, using retrograde trac- ing in the rhesus monkey), suggesting reciprocal con- nections and more complex interactions between these regions. Note that the correspondence between the PAC and STS region found here cannot be explained by overlap between these areas: There was no overlap in five deaf participants (0.8% for the remaining one par- ticipant) and no overlap at the group level in the right hemisphere. The Right STS Region Is Recruited Extensively for Processing Direction-selective Visual Motion in Early Deaf Individuals The most striking difference between deaf and hearing individuals in response to directional motion was found in the right STS region, which was 5.5 times larger for deaf than hearing participants (for a 12 times greater extent in the right posterior STS in deaf than hearing participants in response to attended visual motion, see Table 2 of Bavelier et al., 2001). In contrast, no dif- ferences in direction-selective responses were found across groups in the left STS region or visual area hMT+ here. The STS is a likely region for cross-modal organization, as it covers an expansive region of the temporal lobe and expresses great functional diversity, containing sub- regions sensitive to auditory, visual, tactile, and multi- sensory stimuli (e.g., Dahl, Logothesis, & Kayser, 2009; Beauchamp et al., 2008; Beauchamp, Argall, Bodurka, Duyn, & Martin, 2004; Calvert, Campbell, & Brammer, 2000; Seltzer & Pandya, 1978; Benevento et al., 1977). The posterior STS receives inputs from both the visual and auditory cortex, whereas the middle STS normally receives auditory inputs only, at least in the rhesus mon- key (Seltzer & Pandya, 1994); in humans, auditory–visual responses have been reported to be largest in the middle STS ( Venezia et al., 2017). Congruently, the auditory association cortices have also been invoked in studies in neurotypical individuals on cross-modal plasticity through learned associations (e.g., Meyer, Baumann, Marchina, & Jancke, 2007; see also Bulkin & Groh, 2006; Ghazanfar & Schroeder, 2006). In congenitally deaf people, greater connectivity be- tween the middle STS across hemispheres, as well as with the ipsilateral posterior STS, hints at a reorganization of this region in line with cross-modal plasticity (Li, 2013). Specific examples of cross-modal plasticity in the STS re- gion have been reported with regard to how early deaf people process sign language. Early deaf participants have been shown to have increased activation in the middle STS in response to sign language (e.g., Sadato et al., 2004; Neville et al., 1998). In addition, increased posterior STS activation was shown in deaf signers, and not hearing signers, when performing a velocity task (see Figure 6 of Bavelier et al., 2001). Our report of ex- pansive recruitment of the STS region in early deaf people in response to visual motion thus further con- firms a general pattern of neural plasticity in this region. In hearing individuals, some authors claim that re- sponses to auditory motion are separate from those to auditory localization and rely on the superior temporal gyrus (e.g., Ducommun et al., 2002, 2004; Baumgart, Gaschler-Markefski, Woldorff, Heinze, & Scheich, 1999). However, others claim that the auditory cortex may be selective to spatial locations rather than motion (e.g., Smith, Okada, Saberi, & Hickok, 2004). Our findings sug- gest that, at least in response to visual motion, auditory areas in deaf participants and association areas in deaf and hearing participants are selectively responsive to directional visual motion. Direction-selective Responses to Stimuli in the RVF and LVF Do not Show a Contralateral Bias in the Deaf PAC and Association Cortex Interestingly, despite a right-hemisphere advantage for early deaf participants in the STS region, and hinted at in the PAC, there was more activation overall to stimuli presented in the RVF. Behaviorally, an RVF advantage for visual motion perception has often been reported for early deaf participants (e.g., direction of motion: Neville & Lawson, 1987; direction of motion: Bosworth & Dobkins, 1999; motion velocity: Brozinsky & Bavelier, 2004; direc- tion of motion: Bosworth et al. 2013; see also Samar & Retter, Webster, and Jiang 1135 l D o w n o a d e d f r o m h t t p : / / d i r e c t . m i t . e d u / j / o c n a r t i c e - p d l f / / / 3 1 8 1 1 2 6 1 7 8 8 6 7 7 / / j o c n _ a _ 0 1 3 7 8 p d . f b y g u e s t t o n 0 7 S e p e m b e r 2 0 2 3 Parasnis, 2007; but see Hauthal et al., 2013, for an LVF ad- vantage for movement localization in late signers). A right-hemisphere advantage has been reported in previous neuroimaging studies investigating visual mo- tion or flickering stimulus responses in early deaf people: In the auditory cortices, the right hemisphere was dom- inantly (Finney et al., 2003) or exclusively (Fine et al., 2005; Finney et al., 2001) activated. The right-hemisphere advantage reported here is also in line with the finding that only the right auditory cortex (planum temporale) showed a correlation between increased cortical thick- ness and enhanced visual motion detection thresholds in the study on early deaf people of Shiell et al. (2016). The right auditory cortex was also shown with ERP source localization to be dominant in hearing-restored deaf indi- viduals when viewing visual stimuli (Sandmann et al., 2012). Although some studies have reported left- hemisphere advantages in early deaf people, such effects either could not be localized (e.g., Neville & Lawson, 1987) or were relatively small, sometimes nonsignificant effects, reported in hMT+ (e.g., Fine et al., 2005; Bavelier et al., 2001), which in our study, on direction- selective responses, did not show significant differences across participant groups but an appearance of right lat- eralization for early deaf participants only (see Figures 2B and 4B). Here, we can reconcile behavioral RVF and neural right-hemisphere advantages in response to directional motion: The remapped auditory cortices do not show a strong contralateral bias like other direction-selective areas, for example, hMT+; instead, the right hemisphere dominates regardless. As addressed in the first section of the Discussion, the STS region possesses large spatial fields and, particularly in the right hemisphere, low sen- sitivity to retinotopic organization (e.g., Saygin & Sereno, 2008; see also Almeida et al., 2015; Grossman & Blake, 2001). The recruited association cortex seems to be the best candidate for behavioral RVF advantages, because this region showed the most extensive changes between early deaf and hearing participants here. In addition, in a previous study reporting effects in hMT+, the posterior STS was shown to be 9.3 times larger in size (in compar- ison, hMT+ was only 1.08 times larger) and seven times greater in percent signal change (hMT+: 1.05 times greater) in deaf than hearing participants in response to attended velocity of visual motion (see Table 4 of Bavelier et al., 2001). A right-hemisphere advantage paired with an RVF ad- vantage goes against the assumption that neural activa- tion to visual stimuli is necessarily contralateral, which has frequently been made in the literature on neural plasticity in early deaf people (e.g., Bavelier et al., 2001; tentatively in Bosworth et al., 2013; Hauthal et al., 2013; Brozinsky & Bavelier, 2004; Bosworth & Dobkins, 1999, 2002). In a study reporting a left-hemisphere advantage with attention-related modulation of ERPs to peripheral visual targets in early deaf people, Neville and Lawson (1987) hypothesized that the left hemisphere was re- mapped for sign language processing and therefore could have different sensitivities to stimuli such as visual motion or stimulus localization. However, it is not clear that the left hemisphere is specialized for sign language processing in early deaf people: Deaf and early-signing hearing participants have been shown to have bilateral (STS) activation to sign language; and early deaf partici- pants, to have more right STS activation to written lan- guage (e.g., Sadato et al., 2004; Neville et al., 1998). Our finding of a right-hemisphere advantage compatible with an RVF advantage offers an alternative explanation and unites most neural and behavioral findings regarding motion perception of early deaf people. Speculatively, our findings suggest that behavioral advantages in early deaf people, particularly for motion discrimination in the RVF, may be supported by increased STS region activation in the right hemisphere (again, see Figure 4A). Although limited by a small sample size, five of our six deaf participants also participated in a behavioral study in our laboratory, in which thresholds on the percent dot coherence required for direction of visual motion in the LVF and RVF were acquired. We found a suggestive correlation with this measure of behavioral di- rection discrimination ability (i.e., lower percent dot motion coherences required) and the extent of activation in the right STS region, R2 = .30, although not significant, p = .10. In comparison, the extent of bilateral hMT+ activation showed no correlation, R2 = .01, p = .78. How- ever, this tentative result would need to be confirmed with larger sample sizes in future studies. At the least, we are able to introduce the hypothesis that the auditory and association cortices in early deaf individuals are sensitive to directional visual motion and that this neural reorga- nization may support a behavioral advantage reported previously for visual motion direction discrimination. Limitations and Future Directions One limitation of this study was that the sample con- sisted of only six participants per group: Although the re- sults were reliable across individual participants, they would be strengthened by replication in future studies, potentially with larger sample sizes. Another limitation here is that an eye tracker was not used during the fMRI experiment to ensure fixation. However, it is un- likely that eye movements could explain the results. The functionally defined regions were localized with centrally presented stimuli, and the differences between deaf and hearing participants for peripherally presented stimuli were highly specific (e.g., the enhanced STS re- gion recruitment was restricted to the right hemisphere and RVF). In addition, cues for participants to report the direction of motion were given at random intervals throughout the experiment, such that they were neither periodic nor associated with directional motion presenta- tion times (see Methods). A third limitation of this study 1136 Journal of Cognitive Neuroscience Volume 31, Number 8 l D o w n o a d e d f r o m h t t p : / / d i r e c t . m i t . e d u / j / o c n a r t i c e - p d l f / / / 3 1 8 1 1 2 6 1 7 8 8 6 7 7 / / j o c n _ a _ 0 1 3 7 8 p d . f b y g u e s t t o n 0 7 S e p e m b e r 2 0 2 3 was that the STS region was defined broadly in each participant; future studies could use anatomical land- marks or more specific functional localizers in hearing participants to demarcate more precise subregions. Finally, in this study, directional visual motion coincided with coherent visual motion. Although it may be ar- gued that coherent motion inherently possesses direc- tionality, future studies may address the influence of coherency on directional motion responses (see Braddick et al., 2008). Acknowledgments This research was supported by grants from the National Institutes of Health (NIH; grants EY023268 to F. J. and P20 GM103650 to M. A. W.). 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