Synopsis
Behavioural and fMRI studies have revealed enhanced auditory
abilities and differences in the brain organization of early blind individuals
compared to sighted. We used fMRI to assess cortical interactions associated
with speech processing of repeated and novel stimuli. Although the in-scanner
behavior for the blind and sighted groups were comparable, there were a number
of cortical activation and deactivation differences. Additionally, the blind
participants were better at a post-scan recall task. Cross-modal recruitment of
occipital areas was found in the blind participants. They also showed
widespread repetition-enhancement effects, suggesting that additional attention
mechanisms contribute to their enhanced auditory word encoding abilities.
Introduction
In the past two decades, a great deal of scholarly
literature has established that the functional organization of the brain of early
blind people (EB) differs from that of sighted individuals (SI). As for
perceptual processing, fMRI studies reveal that long-term cross-modal plasticity
is characterized by the recruitment of different cerebral networks. For speech
processing, in addition to auditory areas, visual and audiovisual areas are
more activated in EB 1-5. Differences
in perceptual processing between EB and SI are also observable at the
behavioral level 3,6-10. The EB
constitute a particularly interesting model to understand the developmental plasticity
and reorganization characteristics of processing networks and their behavioral manifestations.
To get a more comprehensive view of the underlying mechanisms associated with
enhanced auditory processing we explored another aspect of cerebral function; neural
adaptation (NA). NA is a phenomenon in which the repetition of a stimulus triggers
a change in the response to the stimulus, upon repeated presentation. This
mechanism is well known in neurophysiology, where these adaptation mechanisms
occur at the level of the single neuron or group of neurons. FMRI adaptation
protocols have highlighted similar mechanisms at the scale of BOLD signal
measurement. Repetition-suppression (RS) is associated with a reduction in the
BOLD response in areas processing a repeated stimulus, and the opposite
response, repetition-enhancement (RE) is associated with stimulus recognition,
learning, attention, expectation and explicit memory 11.
In a previous study 2 RS was observed in auditory areas of both EB and SI during
passive perception of repeated vowel sounds. In EB, we observed additional
areas of RS that included visual (occipital) and audiovisual (superior parietal
lobule) processing regions. In the current study, words and pseudowords were
presented repeatedly. In contrast to our previous study, the repetitions were
presented discontinuously from one another and participants were engaged in an
active lexical decision task.
Methods
Fourteen EB participants and fourteen SI participated in the
study. Functional volumes were acquired using a sparse sampling design to avoid
acoustic scanner noise12. Stimuli were presented either
once or 6 times throughout the experiment. After each trial, participants made
lexical judgments (whether the stimulus item existed or not). After the MRI
session, participants were tested on the material they heard during the MRI
session.
Results and Conclusion
The results of the lexical decision task show that words
were answered faster than pseudowords (F(1,28) = 66.5, p<10^-5), that there
was a repetition priming effect (F(1,28) = 207.2, p<10^-5) and that there
was no significant difference between the groups reaction time (F(1,28) = 0.06,
p>0.8). However, the EB were significantly better to recall if a stimulus had
been presented or not during the experiment (F(1, 28) = 8.22, p=.0078). During stimulus
perception, temporal, parietal and frontal areas traditionally involved in
lexical decision tasks were activated in both groups (Fig 1,2). In addition the
EB showed cross-modal activity in the occipital lobe (Fig 1). Two-sample
t-tests showed no significant difference between the groups for words. However,
for pseudowords the EB activated significantly more than SI, portions of the
precuneus, cuneus, fusiform gyrus, inferior occipital gyri, middle occipital
gyri and superior occipital gyri. The SI had significantly more activity than EB
in bilateral clusters around the hippocampus, as well as in the insula and
Heschl’s gyrus for unique pseudowords (Fig 3). Regarding adaptation effects,
the controls showed repetition-suppression effects in the left inferior frontal
gyrus, and repetition-enhancement in the right SMG, left precuneus and right
postcentral gyrus (Fig 4). The blind did not show any repetition-suppression
effect, however they showed widespread repetition-enhancement effects in
portions of the parietal lobe and frontal lobe including the angular gyrus, the
precuneus, the pre and post-central gyri, as well as in the middle occipital
gyrus, and left hippocampus (Fig 5). Our first study indicated that
repetition-suppression was occurring during the passive perception of repeated
sounds, and that these mechanisms were also present cross-modally in EB. Here,
we show widespread repetition-enhancement effects during the active processing
of words and pseudowords in EB only. The presence of repetition-enhancement
effects seems to indicate that during an active task, additional phenomenon,
such as stimulus recognition, expectation and more generally attention
mechanisms could contribute to the enhanced ability of blind participants to
process auditory stimuli. Taken together the results indicate that the
long-term neuroplastic reorganization occurring in the blind brain not only
involves cross-modal plasticity of sensory areas but also a functional
reorganization of auditory attention mechanisms that contribute to the enhanced
auditory encoding of sounds.
Acknowledgements
Research supported by a Discovery Grant from NSERC (awarded
to V. Gracco).
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