Anissa L. Ramadhani1,2, Ali-Reza Mohammadi-Nejad1,2,3, Katrin Krumbholz2,4, and Dorothee Auer1,2,3
1Radiological Sciences, DCN, School of Medicine, University of Nottingham, Nottingham, United Kingdom, 2SPMIC, School of Medicine, University of Nottingham, Nottingham, United Kingdom, 3National Institute for Health Research (NIHR), Nottingham BRC, Queens Medical Ctr, Nottingham, United Kingdom, 4Hearing Sciences, DCN, School of Medicine, University of Nottingham, Nottingham, United Kingdom
Synopsis
Functional magnetic resonance imaging (fMRI) studies have
suggested that neural variability and connectivity (temporal coherence) are
generally reduced during active versus passive (or resting) brain states. In
this study, we investigate if this reduction in brain coherence is a
supra-modal or modality-specific effect. For that, we compared BOLD signal
during rest and whilst performing a continuous auditory or visual task. Our
results suggest that task-induced reduction in brain coherence occurs only in
default-mode and hetero-modal brain regions.
Introduction
During rest, brain regions
activate coherently at both local and global (long-range network) levels. In
contrast, in the active state, when performing a task or receiving a sensory
stimulus, brain coherence has been shown to be reduced, prompting the
suggestion that decreased brain coherence is associated with increased
information capacity1,2,3. However, it is unclear whether reduction in brain coherence is
specific to brain regions responding to the external stimuli or task (henceforth referred to as homo-modal
regions), or whether it is more widely affecting
long-range networks and
occuring also in brain regions not associated with the task or stimulus
modality (referred to as hetero-modal regions). Evidence for the latter has
come from studies of cross-modal deactivation4,5. Understanding the detailed relationship between task
conditions and brain coherence may help us to understand how phantom
perceptions such as tinnitus are generated and maintained and how they, in turn,
may affect supra-modal brain processes.
We used BOLD fMRI to compare
spontaneous activity and functional connectivity changes during an auditory and
a visual task and during rest. Both tasks required continuous task engagement
throughout an imaging run.Methods
Data acquisition
A total of 11 healthy participants (3 female, mean age = 29
years, age range = 18-35 years) took part in this study. Imaging was performed
on a 3T GE Discovery scanner with 32-channel head coil (Sir Peter Mansfield
Imaging Centre, Nottingham). Functional images were acquired using a
gradient-echo EPI sequence (TR = 2000 ms, TE = 32 ms, voxel resolution = 3x3x3
mm3, 0.5 mm inter-slice gap). We acquired three fMRI scans
consisting of 240 volumes each (two task scans and one resting-state scan). In
addition, we also acquired a T1-weighted structural image using a Sagittal
FSPGR Bravo sequence (TR = 8.156 ms, TE
=3.172 ms, resolution = 1x1x1-mm3, FOV = 256 × 256 × 156).
Task conditions
During the task scans,
participants either watched a silent movie or listened to a radio podcast,
presented at a comfortable listening level. The movie and podcast were
presented through MR-compatible goggles (VisuaStim Digital) or headphones
(SereneSound; both Resonance Technology Company, USA). Participants were told
to close their eyes for the resting-state and auditory task scans, and to stay
awake during the resting-state scan. After scanning, participants were asked
about their experience of the movie and podcast.
Data processing and analysis
Structural and functional data
were processed using the BRC pipeline (v1.4.3)6 in FSL (v6.0.1). The
pre-processing included brain
extraction, motion correction, slice-timing correction, spatial smoothing (5-mm
FWHM Gaussian kernel), ICA-AROMA, and conversion to standard space (2x2x2 mm3). The pre-processed data were then further denoised by means of
scrubbing (FD > 0.5 mm or DVARS > 3%)7, and band-pass
filtering between 0.01-0.08 Hz. Regional connectivity density, interregional
correlations and fractional amplitude of low-frequency fluctuations (fALFF)
were calculated using the CONN functional connectivity toolbox (v19c). Auditory
cortical region of interests (ROIs) were defined using the Brodmann anatomical
atlas. Inter-regional correlations were Fisher z-transformed.Results
Functional connectivity during
rest was largely limited to regions of what is typically know as the
default-mode network. During both task conditions (auditory and visual),
functional connectivity within these regions was reduced, but connectivity was
increased in modality-specific regions—specifically, regions associated with
the languange network in the auditory task and with the dorsal-visual network
in the visual task.
During rest, the auditory
cortical regions (Fig. 2A) showed particularly strong connections between
primary and secondary regions intra-hemispherically, and between the left
secondary and tertiary regions, which are thought to be involved in speech
perception8 (Fig. 2B). Generally, the visual task caused small but
consistent reductions in connectivity strength across all auditory regions. In
contrast, the effects of the auditory task were less consistent, with the
intrahemispheric connections generally increasing in strength, but the
interhemispheric connection generally decreasing in strength (with the
exception of the interhemispheric connection between the primary regions; Fig
2C).
The local variability
(represented by fALFF), showed a consistent pattern of regional differences,
with the primary region showing less variability than the secondary and
tertiary regions. During the auditory task, variability across all three
regions tended to either remain unchanged or be slightly increased. In contrast,
during the visual task, variability was consistently decreased across all three
regions.Discussion and conclusion
Contrary to the
idea that performing a task causes general reduction in brain coherence, our
results suggest that coherence reduction is specific to task modality,
occurring only, or predominantly, within default-mode and hetero-modal regions,
with homo-modal regions showing either
no change or even increase in coherence. These results are consistent with the
idea that performing a task increases long-range inhibitory drive exerted by
the task-activated (homo-modal) regions, whilst, at the same time, reducing
local inhibitory drive within these regions9. These, albeit
preliminary, challenge the presumed association between brain coherence and
information capacity. Instead, they suggest that increased brain coherence may
actually facilitate information processing, perhaps through modulation
of neural excitability at finer temporal or spatial scales.
Assuming that the auditory task
condition can be taken as a model of tinnitus, our results suggest that
tinnitus should increase brain coherence in auditory-specific brain regions but
decrease coherence in the default mode network and in non-auditory sensory
networks.Acknowledgements
*Author KK and DA share senior authorship.
This research is a part of the TINACT
project and has received funding from the European Research Council (ERC) under
the European Union's Horizon 2020 research and innovation programme (grant
n°764604). We thank the NIHR Nottingham Biomedical Research Centre and
Precision Imaging Beacon for valuable intellectual input.
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