Evgeniya Kirilina1,2, Ilona Lipp1, Carsten Jäger1, Markus Morawski3, Merve N. Terzi4,5, Hans-Jürgen Bidmon6, Markus Axer4, Pitter F. Huesgen5, and Nikolaus Weiskopf1,7,8
1Department of Neurophysics, Max Planck Institute for Human Cognitive and Brain Sciences, Leipzig, Germany, 2Center for Computational Neuroscience, Free University Berlin, Berlin, Germany, 3Paul Flechsig Institute of Brain Research, University of Leipzig, Leipzig, Germany, 4Institute of Neuroscience and Medicine, Forschungszentrum Jülich, Juelich, Germany, 5Zentralinstitut für Engineering, Elektronik und Analytik, Forschungszentrum Jülich, Juelich, Germany, 6C. und O. Vogt-Instituts für Hirnforschung, Heinrich-Heine-Universität Düsseldorf, Duesseldorf, Germany, 7Felix Bloch Institute for Solid State Physics, Leipzig University, Leipzig, Germany, 8Wellcome Centre for Human Neuroimaging, University College London, London, United Kingdom
Synopsis
Quantitative MRI parameters in the brain provide
unique information on tissue myelination. However, the validation studies performing quantitative comparisons between
MRI metrics and tissue myelin content are very limited, mainly due to the to
the lack of methods for histological myelin quantification. Here, we explore
lipid imaging using matrix-assisted laser desorption/ionization (MALDI) and
multiple histological myelin stains in post-mortem human brain tissue samples for
validation of MRI based myelin biomarkers. We show that tissue lipid
composition vary across different cortical layers and white matter pathways,
potentially reflecting differences in myelin structure and may impact MRI-based
myelination metrics.
INTRODUCTION
Myelin, the fatty axon-insulating
substance in the brain, is composed of a large variety of lipids, proteins and
trapped water1. It is the main source of contrast in magnetic
resonance images (MRI) of the human brain1–3. All MRI parameters, including longitudinal and
transverse relaxation rates (R1, R2, R2*), proton density (PD), magnetisation
transfer and magnetic susceptibility are sensitive to tissue myelination due to
different biophysical mechanisms1,3–6. Therefore, quantitative MR parameters used as
myelination biomarkers provide unique in
vivo information on brain development, cortical myeloarchitecture,
plasticity and neurodegeneration7–9.
However, the mechanisms
underlying the sensitivity of MRI parameters to tissue myelination are only
partly understood and quantitative comparisons between MRI metrics and tissue
myelination are limited to some few studies1,10. Moreover,
specificity and sensitivity of R1, R2* and PD to myelin composition are only starting
to be explored10. Validation of MRI-based myelin biomarkers is
difficult due to the lack of methods for histological myelin quantification.
Classical histological and immunohistochemistry stains provide only qualitative
information on myelin distribution and reflect only some of the various myelin
components. Recently developed advanced methods for lipid quantification promise
to overcome this limitation11.
Here, we systematically
explore several methods for myelin quantification for the validation of MRI
myelin biomarkers. To this end, we combined quantitative MRI in post mortem human brain tissue samples
with histological and immunohistochemical myelin stainings and lipid imaging
with matrix-assisted laser desorption/ionization mass spectrometry (MALDI-MSI).METHODS
We investigated tissue blocks from five human post-mortem human brains, all
containing the primary and higher visual cortices12. Multi-parametric maps13, including quantitative maps of R1, R2* and proton
density (PD), were obtained on a 7T MRI system (Magnetom, Siemens Healthineers,
Germany), using two FLASH acquisitions (TR=180ms, TE1-16=2.4-40ms, FA: 12° and
65°, isotropic resolution of 0.2 mm, additional calibration scan for flip angle
(FA) mapping) and a customized version of hMRI toolbox (hMRI.info).
Macromolecular volume fraction (MVF) maps were calculated using PD maps3.
After completing MRI scanning, tissue blocks were infiltrated with 30%
sucrose and 30 µm thick cryosections were obtained. Consecutive sections were
stained with several myelin stains: a modified Gallyas stain14 (silver impregnation), Luxol Fast Blue15 (histological stain) and an antibody directed against
myelin basic protein (MBP immunohistochemistry). Additionally, two of the
samples underwent lipid assessment with MALDI-LTQ-Orbitrap (Thermo Scientific
Fisher, Schwerte) using mass range of 400-1000 m/z. For the MALDI-MSI data, a
principal component analysis (PCA) on 65 mass peaks corresponding to known
lipids was computed. We then co-registered the MALDI-MSI lipid distribution
maps with the quantitative MRI maps and performed voxel-wise correlation
analysis.RESULTS AND DISCUSSION
All myelin stains and all quantitative MR parameters showed high contrast
between white and grey matter, with white matter having higher myelin content.
However, the patterns of myelination revealed in the cortex differed between stains
and parameters (Fig. 1). A gradient of myelin content from the pial surface to
white matter boundary was prominent in the silver stain and anti-MBP immunohistochemistry,
resembling the patterns observed in R2* and PD maps. Hyperintensity of layer IV
was observed in the Luxol Fast Blue staining, corresponding to the pattern
found in the R1 maps.
MALDI-MSI revealed distinct distributions of lipid components in white and
grey matter, cortical layers and in optic radiation (Fig. 2). This is important
for interpretation of R1- and MT-based myelin biomarkers, since it has recently
been demonstrated that lipid composition can influence relaxation properties of
myelinated fibers10. The PCA suggested the presence of two main lipid
components. Lipids of the first component were predominantly located in white
matter, also demonstrating an intra-cortical gradient, and most likely reflect
local myelin content (Fig.2). The lipids of the second components were mostly
localized in the primary visual cortex. High spatial correlations between the
first PCA component and the MRI metrics were observed (pixel-wise Pearson
correlation coefficient of r=0.66 for
R1, r=0.78 for R2* and r=0.84 for MVF), with the proton-density based measure
showing the highest correlation with the abundance of myelin lipids.CONCLUSIONS
We demonstrate that different histological stains for myelin detection
provide similar, but distinct information on tissue myelination, particularly
in the cortex. Differences between histological methods potentially correspond
to differences in specificity between MRI-based myelin markers. Therefore, the assessment of cortical
myelin with histology is not
comprehensive and requires the use of multiple stains. We show that
lipid composition of tissue varies across different cortical layers and white
matter pathways, potentially reflecting differences in myelin structure. We
suggest that a principal component analysis of MALDI-MSI lipid maps can be used
to obtain a histological myelin biomarker for the validation of quantitative
MRI parameters. Our results demonstrate that MALDI-MSI is a powerful tool for
validation of myelin MR markers and that difference in lipid composition of the
cortex and in white matter pathways should be taken into account when
interpreting MRI-based maps of brain myelination.Acknowledgements
The research leading
to these results has received funding from the European Research Council under
the European Union's Seventh Framework Programme (FP7/2007-2013) / ERC grant
agreement n° 616905 and the BMBF (01EW1711A & B) in the framework of
ERA-NET NEURON. Further, this work was supported by the German Research
Foundation (DFG Priority Program 2041 ”Computational Connectomics”, MO 2249/3-1
and the Alzheimer-Forschung-Inititiative e.V., (AFI # 18072).References
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