Helene Benveniste1, Gerald Dienel2, Zvi Jacob3, Hedok Lee1, Rany Makaryus3, Albert Gjedde4, Fahmeed Hyder5, and Douglas L. Rothman5
1Anesthesiology, Yale School of Medicine, New Haven, CT, United States, 2Department of Neurology, University of Arkansas for Medical Sciences, Little Rock, AR, United States, 3Anesthesiology, Stony Brook Medicine, Stony Brook, NY, United States, 4Panum Institute, University of Copenhagen, Copenhagen, Denmark, 5Department of Biomedical Engineering & Radiology and Biomedical Imaging, Yale School of Medicine, New Haven, CT, United States
Synopsis
Lactate is produced in normal brain even when
O2 levels are ample, and high lactate
production may explain aerobic glycolysis (AG) in developing brain. We evaluated steady
state lactate concentrations ([Lac]) in brains of 87 children using 1HMRS while they underwent
routine MRI examination. The trajectory of [Lac] in cerebral cortex
across childhood was below the 0.5-0.7 mM range in normal adult brain. Thus, lactate accumulation and efflux
are unlikely to underlie excessive AG in children.
Introduction
Characterizing
the metabolic needs of the developing brain is essential for brain health and
for understanding disease processes arising during childhood and adolescence. The
growth pattern of brain development is complex and is paralleled by age-varying
energy requirements. A disproportionately higher rate of glucose utilization
(CMRglc) ( ̴33%) compared
with oxygen consumption (i.e. aerobic glycolysis, AG) was documented in
children’s brain and proposed to support developmental processes1, 2. Several candidate metabolic pathways may
explain the non-oxidative glucose consumption and lactate production is
considered a major contender. The ≈33% excess CMRglc equals 0.18 μmol
glucose/g/min or 0.36 μmol/g/min lactate release from brain. Here we characterize the trajectory of the cortical
lactate concentration, [Lac], across childhood to determine if indeed lactate
accumulates in brain and is high enough to account for the glucose consumed in
excess of oxygen and to support a high rate of lactate
efflux from the young brain.Methods
De-identified 1HMRS spectra and anatomical
T1-weighted scans from 87 children undergoing routine MRI were used. 1HMRS
from 60 of the 87 children were previously reported in a study focusing on
another topic3. The 1HMRS spectra were analyzed using LCModel. To assess the magnitude of
glucose-oxygen uncoupling consistent with the measured lactate levels we also
calculated the cerebral metabolic rate of lactate, CMRlac according
to Eq. 1-3.
$$ CMR_{Lac} = V_{in} - V_{out}\quad\quad\quad\quad\quad(Eq. 1)$$
$$
V_{in} = V_{MAX}\frac{[Lac]_{p}}{K_{T}+[Lac]_{p}+[Lac]_{B}}\quad\quad(Eq. 2)$$
$$
V_{in} = V_{MAX}\frac{[Lac]_{B}}{K_{T}+[Lac]_{p}+[Lac]_{B}}\quad\quad(Eq. 3)$$
Where [Lac]p and [Lac]B are the
concentrations of lactate in plasma and brain, respectively. Since we did not
measure [Lac]p in the children we set it to be 0, 1, or 2 mM for the
calculations, according to normal clinical reference ranges. Lactate transport
kinetic parameters VMAX and KT determined previously in
adult brain of 0.4μmol/g/min
and 5.1 mM4,
respectively, were used for the calculations.
Results
Nine 1HMR spectra were excluded from the
analysis due to poor SNR and wild baseline fluctuations; leaving 78 for analysis.
The average FWHM and SNR of the spectral NAA peaks was 0.028 ± 0.006 ppm and
22.3 ± 4.2, respectively. Fig. 1
shows typical cortical 1HMRS spectra from a 3- and a 7-yr old child.
The Cramer–Rao lower bounds (CRLBs) calculated by LCModel for [tNAA] were 2-5
%SD. The CRLB’s for [Lac] were considerably higher; and [Lac] values associated
with CRLB’s >350% were discarded; leaving 71 subjects with [Lac] for
analysis. The high CRLB for [Lac] was due to its low concentration being on the
order of the noise level in some subjects; however the %SD variation was at most 0.2 mM as
estimated from the CRLB of the NAA peak. Volumetric changes in
total gray and white matter were positively correlated with age (Fig. 2). Peak [Lac] occurred at ≈5 years of age (Fig. 3), was lower than predicted, and below the range for adult
brain (0.5-0.7mM)5. The calculated CMRlac
indicated a net influx of +0.04 μmol/g/min. In other words, the
prediction of lactate release from the developing brain (CMRlac = -0.36 μmol/g/min) was not substantiated.Discussion
In contrast to reports of high AG1
predicting a high rate of lactate efflux from the young brain we found that
brain [Lac] was low in children across childhood and in the range or lower than
reported in adult subjects. Further, the [Lac] corresponded to a net lactate
influx of +0.04 μmol/g/min.
The ability to measure resting lactate by 1HMRS has been criticized
based on its low levels and contamination from brain macromolecules and lipids
from the skull. However, if all the resonance intensity at 1.3 ppm was due to
lactate, its concentration would be at most ~1 mM (Fig. 1), which is still well below what would be needed to explain
the reported oxygen-glucose mismatch.
The oxygen-glucose index (OGI) is the metric from which AG
is quantified and the theoretical OGI is 6.0. The inference is that glycolytic
flux is increased (and OGI decreased) in the developing brain but the
downstream fate of the glucose carbon is not established. For example, an
alternate possibility to explain the elevated AG is the pentose phosphate
pathway. Furthermore, OGI will
exceed 6 if supplemental substrates (β-hydroxybutyrate,
lactate) are oxidized but not measured and not included in the OGI calculation
with glucose.Interpretation
Accumulation of [Lac] and calculated efflux of
lactate from brain do not explain AG. However, the value for the lumped
constant for [18F]fluorodeoxyglucose has a high impact on calculated
CMRglc. Use of updated values for the lumped constant alters or eliminates
the oxygen-glucose mismatch in developing brain and will be presented.Acknowledgements
DLR - 1- R01NS087568A, R01NS100106; HB - R21HD080573; FH - R01MH067528, P30NS052519References
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