Emmanuelle Flatt1, Bernard Lanz1, Thanh Phong Lê1,2, Rolf Gruetter1, and Mor Mishkovsky1
1Laboratory for Functional and Metabolic Imaging (LIFMET), EPFL, Lausanne, Switzerland, 2Geneva School of Health Sciences, University of Applied Sciences and Arts Western Switzerland (HES-SO), Geneva, Switzerland
Synopsis
The present work describes a quantitative
analysis of lactate 13C-labeling pattern following the metabolism of hyperpolarized
[2H7,U-13C6]-D-glucose. This lactate production results from
12 biochemical steps including glucose transport, 10 enzymatic steps of
glycolysis, and LDH mediated pyruvate-to-lactate conversion. The 3-compartment
model description was found a good compromise to interpret the hyperpolarized
metabolic curves. This demonstrates the potential of hyperpolarized [2H7,
U-13C6]-D-glucose as a new biochemical probe for brain
energy metabolism.
Introduction:
Recent studies have demonstrated the possibility to
monitor lactate production shortly after injection of hyperpolarized [2H7,13C6]-D-glucose
in naïve brain1,2 and brain tumors3. The lactate production from hyperpolarized
13C-glucose results from 12 biochemical steps including glucose transport, 10
enzymatic steps of glycolysis, and LDH mediated pyruvate-to-lactate conversion.
In the present work, we describe kinetic models reflecting the dynamic 13C-labeling
pattern following the metabolism of [2H7,U-13C6]-D-glucose.
Since we detected the dynamics of glucose and lactate solely, we chose to
restrict ourselves to a 3- or 4-compartment model, to avoid the risk of
overfitting (Fig.1). Quantifying the kinetics of cerebral lactate production following
hyperpolarized [2H7,U-13C6]-D-glucose
bolus is an important step to extend the potential of hyperpolarized glucose as
a complement to FDG-PET and thermally polarized X-nuclei MRS
approaches5–8.Methods:
Hyperpolarization: A frozen mixture of 2M [1,2,3,4,5,6,6′-
2H
7,U-
13C
6]D-glucose,
25mM OX63 radical doped with 1mM of gadolinium in water/glycerol
4 was hyperpolarized in a custom-designed 7T/1K DNP polarizer
9, resulting in a liquid-state
polarization of 29±3%.
Animal Preparation: 12 hour-food deprived C57BL6/J male mice (17±2
weeks) were anesthetized using 1.5-2% isoflurane in 60% O
2. The
anesthetized animals were cannulated to place a femoral vein catheter to
deliver the glucose bolus. Post-surgery, a subcutaneous bolus of medetomidine
0.3mg/kg was administered. Ten minutes later, isoflurane anesthesia
was dropped to 0.25–0.5%, and a continuous subcutaneous infusion of medetomidine
(0.6 mg/kg/h) started, as described elsewhere
10.
Acquisition: Mice were placed into a 9.4T/31cm horizontal bore MRI
scanner (Varian/Magnex) with a home-built
1H quadrature/
13C single-loop coil above the head. At 1h post-medetomidine, 540 μL of 44±10mM HP [
2H
7,U-
13C
6]-D-glucose was
injected and
13C MR spectra were acquired every 1s with nominal 25° flip-angle for the C1
lactate resonance (183.5 ppm) and 1.5° flip angle for the glucose C1 resonances
(92.9 ppm and 96.8 ppm).
Kinetic modeling: Spectra were integrated with VNMRJ software to obtain the time course of glucose
and lactate signals. From a simplified scheme of [
2H
7,U-
13C
6]glucose
cerebral metabolism (Fig.1), kinetic
models (Fig.2) were derived as follows:
- Steps
were modeled as first-order reactions.
-
Single
apparent rate constants included glucose transport through the blood-brain
barrier, cell transport, and either glucose-to-pyruvate (3-compartment model) or
glucose-to-intermediate pool (4-compartment model) were assumed.
- Pyruvate-to-lactate exchange is
much faster than the entry into the TCA cycle and the conversion to alanine11, thus, given the short
total acquisition time, neither VMPC nor VALT of pyruvate
were included in either kinetic model.
·
- The rate constant kGlc-Pyr
was forced to be lower than kPyr-Lac (3-compartment model) and
kGlc-IP or kIP-Pyr to be lower than kPyr-Lac (4-compartment
model)11.
- Lactate
and pyruvate resonances are within the RF pulse bandwidth thus the same flip angle
was assumed for both metabolites.
- Signal decay is a combination of the effects of
T1-relaxation, repeated RF excitations, and biochemical
conversions.
Kinetic rate constants were determined
by fitting the models to average metabolites time courses using
Levenberg-Marquart algorithm
12. Monte-Carlo simulations tested
the precision and accuracy of the model. Additionally, for the 3-compartment model, individual animal
time courses were fitted, and we tested the impact of varying both spin-lattice relaxation and flip-angle
parameters.
Results:
Both models could fit the experimental data (Fig.3A).
A better fit was achieved with the 4-compartment model (15% SSE reduction), as
visible in the build-up of the lactate curve.
In the 3-compartment model, both the variation of
the initial conditions and the Monte-Carlo simulations resulted
in kinetic rates within the same range. In the 4-compartment model, this variance was slightly
higher (Fig.3B&C).
Analyzing correlations between the kinetic rate
constants: off-diagonal covariance was increased in the 4-compartment model. In
the 3-compartment model, only kGlc-Pyr and kPyr-Lac are
anti-correlated (Fig.3D).
Testing the robustness of the 3-compartment model: results computed for individual mice were similar to
those obtained from the average time courses confirming the reproducibility of the
data (Fig.4A).
Flip angle variations had a greater influence on the
kinetic rates than the spin-lattice relaxation (Fig.4B&C).Discussion:
Quantitative
information on the kinetics of cerebral hyperpolarized-13C-glucose metabolism
was obtained for the first time.
The
increased variance in kinetic rates and their higher covariance indicate that
the 4-compartment model is probably too complex. Indeed, the two added kinetic
rates (kGlc-IP and kIP-Pyr) were fully correlated and
thus hardly biochemically interpretable. This is
explained by the delay between glucose and lactate signals which is necessarily
created by kGlc-IP or kIP-Pyr steps. Therefore, the rest of our analysis focused
on the 3-compartment model.
The
3-compartment model appeared to be more robust: variation
of initial conditions and Monte-Carlo simulations gave kinetic rate constants
in the same range. Similar values were estimated on average time courses and individual
animal analyses.
The estimated
kPyr-Lac in the 3-compartment model is in the same
range as values reported on isoflurane-anesthetized13
and awake rats14.
Note that the differences may be related to the hyperpolarized-sensor uptake, the anesthesia, and the different
animal species. Conclusion:
Kinetics of cerebral lactate production following hyperpolarized
[2H7,U-13C6]-D-glucose injection was
quantified, enabling hyperpolarized [2H7,U-13C6]-D-glucose
studies to provide biochemical insight on cerebral metabolism and complement existing bioimaging tools.Acknowledgements
We thank Dr. Elise Vinckenbosch for the fruitful discussion on metabolic modeling, and Drs Mario Lepore, Analina Haussin, and Stefanita Mitrea for
providing veterinary assistance for this study.References
1. Mishkovsky M,
Anderson B, Karlsson M, et al. Measuring glucose cerebral metabolism in the
healthy mouse using hyperpolarized 13C magnetic resonance. Sci Rep.
2017;7:11719. doi:10.1038/s41598-017-12086-z
2. Flatt E, Lanz B,
Pilloud Y, et al. Measuring Glycolytic Activity with
Hyperpolarized [2H7, U-13C6] D-Glucose in the Naive Mouse Brain under Different
Anesthetic Conditions. Metabolites. 2021;11(7):413.
doi:10.3390/metabo11070413
3. Mishkovsky
M, Gusyatiner O, Lanz B, et al. Hyperpolarized 13C-glucose magnetic resonance
highlights reduced aerobic glycolysis in vivo in infiltrative glioblastoma. Sci
Rep. 2021;11:5771. doi:10.1038/s41598-021-85339-7
4. Capozzi
A, Patel S, Wenckebach WT, Karlsson M, Lerche MH, Ardenkjær-Larsen JH.
Gadolinium Effect at High-Magnetic-Field DNP: 70% 13C Polarization of [U-13C]
Glucose Using Trityl. J Phys Chem Lett. 2019;10(12):3420-3425.
doi:10.1021/acs.jpclett.9b01306
5. Gruetter R, Novotny
EJ, Boulware SD, et al. Direct measurement
of brain glucose concentrations in humans by 13C NMR spectroscopy. Proc Natl
Acad Sci U S A. 1992;89(3):1109-1112.
6. Lu
M, Zhu XH, Zhang Y, Mateescu G, Chen W. Quantitative assessment of brain
glucose metabolic rates using in vivo deuterium magnetic resonance
spectroscopy. J Cereb Blood Flow Metab. 2017;37(11):3518-3530.
doi:10.1177/0271678X17706444
7. Feyter
HMD, Behar KL, Corbin ZA, et al. Deuterium metabolic imaging (DMI) for
MRI-based 3D mapping of metabolism in vivo. Science Advances. 2018;4(8):eaat7314.
doi:10.1126/sciadv.aat7314
8. Lai M, Lanz B,
Poitry-Yamate C, et al. In vivo 13C
MRS in the mouse brain at 14.1 Tesla and metabolic flux quantification under
infusion of [1,6-13C2]glucose. J Cereb Blood Flow Metab.
2018;38(10):1701-1714. doi:10.1177/0271678X17734101
9. Cheng
T, Capozzi A, Takado Y, Balzan R, Comment A. Over 35% liquid-state 13 C
polarization obtained via dissolution dynamic nuclear polarization at 7 T and 1
K using ubiquitous nitroxyl radicals. Physical Chemistry Chemical Physics.
2013;15(48):20819-20822. doi:10.1039/C3CP53022A
10. Multi-slice
passband bSSFP for human and rodent fMRI at ultra-high field - ScienceDirect.
Accessed November 9, 2021.
https://www.sciencedirect.com/science/article/pii/S1090780719301004?via%3Dihub
11. Quek
LE, Liu M, Joshi S, Turner N. Fast exchange fluxes around the pyruvate node: a
leaky cell model to explain the gain and loss of unlabelled and labelled
metabolites in a tracer experiment. Cancer Metab. 2016;4.
doi:10.1186/s40170-016-0153-9
12. Marquardt
DW. An Algorithm for Least-Squares Estimation of Nonlinear Parameters. Journal
of the Society for Industrial and Applied Mathematics. 1963;11(2):431-441.
13. Marjańska
M, Shestov AA, Deelchand DK, Kittelson E, Henry PG. Brain metabolism under different
anesthetic conditions using hyperpolarized [1-13C]pyruvate and [2-13C]pyruvate.
NMR in Biomedicine. 2018;31(12):e4012. doi:10.1002/nbm.4012
14. Hyppönen
V, Stenroos P, Nivajärvi R, et al. Metabolism of hyperpolarised [1–13C]pyruvate
in awake and anaesthetised rat brains. NMR in Biomedicine.
n/a(n/a):e4635. doi:10.1002/nbm.4635