Laura D Lewis1,2, Kawin Setsompop1,3, Bruce Rosen1,4, and Jonathan R Polimeni1,4
1Athinoula A. Martinos Center for Biomedical Imaging, Massachusetts General Hospital, Boston, MA, United States, 2Society of Fellows, Harvard University, Cambridge, MA, United States, 3Radiology, Harvard Medical School, boston, MA, United States, 4Radiology, Harvard Medical School, Boston, MA, United States
Synopsis
No
current technique can noninvasively localize neural activity in human
subcortical structures at subsecond temporal resolution. Recent studies have
demonstrated that fast (>0.2 Hz) fMRI responses can be detected in human
cortex. We aimed to test whether fast fMRI signals can also be detected in the
thalamus. We presented oscillating visual stimuli in order to induce
oscillatory neural activity in visual thalamus, and observed large-amplitude fMRI
oscillations at 0.5 Hz. We conclude that high-frequency fMRI responses can be
detected in thalamus, suggesting fast fMRI has the potential to be used for
whole-brain imaging.
Introduction
A key strength of fMRI is its ability to image the
whole brain, including subcortical structures such as thalamus that are
challenging to access through other techniques. This imaging is typically performed
at low sampling rates, as the hemodynamic response is classically assumed to be
slow, but recent studies have demonstrated that surprisingly large >0.2 Hz
signals can be detected in human cortex, both in task-driven [1] and resting state [2-6] contexts. Subcortical fMRI
responses are more challenging to measure, since the small size of subcortical
nuclei and the low signal-to-noise in the center of the brain reduces
detectability. Furthermore, the shape and timing of the hemodynamic response
varies substantially over the brain [7-10], and whether fast hemodynamics can
occur in deep structures is not known. We aimed to test whether high-frequency
fMRI signals can be detected in human thalamus. We studied this using
oscillatory visual stimuli to drive oscillating neural activity in primary
visual cortex (V1) and the lateral geniculate nucleus (LGN, a visual nucleus of
the thalamus).Methods
Five subjects gave informed consent and were scanned
on a 7T Siemens scanner with a custom-built 32-channel head coil array. Each
session began with a 0.75 mm isotropic multi-echo MPRAGE [11] and initial locations for V1 and
LGN were identified using anatomical landmarks. Functional runs consisted of 15
oblique slices that were positioned to include both LGN and V1, acquired as
single-shot gradient-echo blipped-CAIPI SMS-EPI [12] with 2 mm isotropic resolution (R=2 acceleration, MultiBand factor=3,
matrix=120×120, CAIPI shift=FOV/3, TR=227 ms, TE=24 ms, echo-spacing=0.59 ms,
flip angle=30°). Visual stimuli were presented for four continuous minutes, in
which a flickering radial checkerboard inverted at 12 Hz. The luminance
contrast of the checkerboard oscillated at either 0.2 Hz or 0.5 Hz in each run
(Fig. 1a). Subjects performed a simple detection task to maintain fixation
throughout the run. For analysis, regions of interest (ROIs) in V1 and LGN were
defined using functional maps from an initial localizer run using 0.1 Hz
stimulation (Fig. 1b), with an anatomical constraint. The frequency response (FR)
was calculated as the ratio of the amplitude of the 0.5 Hz response to the 0.2
Hz response, times 100. Oscillation amplitude was calculated as the magnitude
of the best-fit sine wave. 95% confidence intervals (CI) for the oscillation
amplitude were calculated via bootstrap, resampling subjects and cycles 1000
times.Results
We
first analyzed mean fMRI responses to the 0.2 Hz stimulus, and observed robust
fMRI oscillations both in V1 and LGN (Fig. 2a, V1 amplitude=1.26%, CI=[0.95
1.61]; LGN amplitude: 1.04%, CI=[0.71 1.38]). Canonical models of the
hemodynamic response would then predict undetectable responses at 0.5 Hz, with
a relative frequency response (FR) of 1.3, which would lead to oscillations with
magnitude <0.02%. However, consistent with prior studies [1], we observed oscillations at 0.5
Hz in V1 that were substantially larger than the canonical prediction (Fig. 2b,
V1 amplitude=0.11%, CI=[0.07 0.16]). In addition, 0.5 Hz oscillations of
similar magnitude were observed in LGN (Fig. 2b, LGN amplitude=0.07%, CI=[0.01
0.13]). These results corresponded to an FR of 6.6 in LGN and 9.4 in V1,
several times larger than the 1.3 FR of canonical models. We further observed
that the response in LGN consistently preceded that of V1 by hundreds of
milliseconds (peak 0.2 Hz timing: 1.18 s in V1, 0.82 s in LGN; 0.5 Hz timing:
1.48 s in V1, 1.10 s in LGN). Plotting voxel-level temporal delays demonstrated
that the population of LGN voxels typically responded earlier than V1 by
hundreds of milliseconds (Fig. 3a). In some individual subjects, these temporal
delay distributions were non-overlapping across V1 and LGN (Fig. 3b),
suggesting that hemodynamic responses consistently occur with an earlier onset
in LGN than in V1. Discussion
Our
results demonstrate that >0.2 Hz fMRI signals are present and robustly
detected in human visual thalamus (LGN). The relative frequency response was
very similar across V1 and LGN, suggesting similar hemodynamic nonlinearities
exist in both: specifically, fMRI responses to high-frequency stimuli are
larger and faster than predicted by responses to conventional block-design
stimuli. However, the response phase was consistently different across regions,
with LGN oscillations peaking hundreds of milliseconds earlier than V1. This fMRI
response delay is much larger than the expected neural delay (as V1 neural activity
lags LGN by only tens of milliseconds [13]) and is therefore likely due to
faster hemodynamics in LGN. We conclude that fast fMRI can detect
high-frequency signals in human thalamus, and suggest that mapping local
hemodynamic delays may be crucial for analyzing whole-brain fast fMRI data.Acknowledgements
This work was funded by NIH grants
K99-MH111748, R01-EB019437,
and P41-EB015896; and NCRR shared resource instrumentation grants S10-RR023401,
S10-RR023403, and S10-RR020948.
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