Georgios Batsios1, Meryssa Tran1, Anne Marie Gillespie1, Celine Taglang1, Sabrina Ronen1, Joseph Costello2, and Pavithra Viswanath1
1Radiology and Biomedical Imaging, University of California San Francisco, San Francisco, CA, United States, 2Neurological Surgery, University of California San Francisco, San Francisco, CA, United States
Synopsis
Telomerase reverse transcriptase (TERT) expression is
essential for tumor proliferation. TERT also reprograms metabolism by elevating
NADH and upregulating the alanine transporter ASCT2. Here, we exploit these
TERT-associated metabolic alterations for non-invasive imaging in live GBM
cells and orthotopic tumors using hyperpolarized [1-13C]-alanine.
Combined treatment with the TERT inhibitor 6-thio-2’-deoxyguanosine and the
ASCT2 inhibitor V-9302 inhibits GBM proliferation, identifying a novel
therapeutic opportunity. Importantly, lactate production from hyperpolarized
[1-13C]-alanine is an early biomarker of response to treatment in
vivo, prior to the onset of anatomical alterations. Our findings pave the
way for improved therapy and response assessment for GBM patients.
Introduction
Telomere shortening constitutes a barrier to uncontrolled proliferation1. All tumors must find
a mechanism of maintaining telomere length and most, including high-grade
glioblastomas (GBMs) achieve this via reactivation of telomerase reverse
transcriptase (TERT) expression1,2.
Since TERT is silenced in normal cells and drives tumor immortality, TERT is a
biomarker of tumor proliferation1,2. TERT
is also a therapeutic target and the TERT inhibitor 6-thio-2’-deoxyguanosine
(6-thio-dG) is in cancer clinical trials3.
Studies indicate that TERT elevates NADH and upregulates
the alanine transporter ASCT2 in low-grade oligodendrogliomas, providing a
rationale for the use of hyperpolarized [1-13C]-alanine to monitor
TERT expression4. These
studies also point to the potential therapeutic utility of ASCT2 inhibitors
such V-93025 for
GBMs. Therefore, the goal of this study was to determine whether 1) hyperpolarized
[1-13C]-alanine reports on TERT expression in GBMs 2) a combination
of 6-thio-dG and V-9302 inhibits GBM proliferation and 3) hyperpolarized [1-13C]-alanine
provides a readout of treatment response in vivo.Methods
Cell
models: We studied patient-derived GBM models (GBM1 and GBM6) that
use TERT for telomere maintenance. Cells were maintained as described previously6,7.
Hyperpolarized
alanine preparation: [1-13C]-alanine was prepared as
described4,
polarized for ~1.5h, and dissolved in isotonic buffer to a final concentration
of 38.9mM (cell studies) or 195mM (in vivo studies).
Treatment:
siRNAs against TERT or non-targeting siRNA
pool were used to transiently silence gene expression in GBM1
and GBM6 cells4. Alternatively, cells were
treated with vehicle control (saline), 6-thio-dG (10μM), V-9302 (10μM) or a combination (10μM each) for 72h. For in
vivo studies, rats bearing orthotopic GBM6 tumors were intraperitoneally
treated with a combination of 6-thio-dG and V-9302 (50mg/kg each) daily.
Hyperpolarized
13C-MRS in live cells: Hyperpolarized [1-13C]-alanine
was injected into ~3x107 live cells in a 5mm tube and 13C-MR
spectra acquired every 3s for 300s on a Varian 500MHz spectrometer using a 13° pulse. Signal-to-noise (SNR)
ratios were quantified in summed spectra using MestReNova.
Hyperpolarized
13C-MRS in vivo: MR studies were performed on
a horizontal Bruker 3T scanner equipped with a quadrature 1H-13C
volume coil. Athymic nu/nu rats were intracranially injected with 3x105
GBM6 cells4,8. Tumor
growth was assessed by T2-weighted MRI using a spin echo (TurboRARE) sequence4,8. Following
intravenous injection of 2.2ml of hyperpolarized [1-13C]-alanine, 2D
echo planar spectroscopic imaging (EPSI) data was acquired with a spatial resolution
of 5.375x5.375x8mm3 (TR=3s)4,8. 13C
spectra were analyzed by calculating the area under alanine and lactate peaks.
Intensity heat maps were produced by interpolating the data using a Lanczos-2
kernel. Ratios of lactate/alanine SNR were evaluated in Matlab4,8.
Statistical analysis: All
results are expressed as mean±STD. Statistical significance was assessed using
an unpaired two-tailed Student’s t-test with p<0.05 considered significant.Results and Discussion
Lactate production from hyperpolarized [1-13C]-alanine
is a biomarker of TERT expression in GBM cells: Fig
1A-1B show a representative 13C spectral array and summed 13C
spectra from GBM1 TERT+ and TERT- cells. Importantly, lactate production from
hyperpolarized [1-13C]-alanine was significantly reduced in TERT-
cells relative to TERT+ in both GBM1 (98.6% drop, p=0.006; Fig.1C) and GBM6 (98.3%
drop, p=0.008; Fig.1C) models.
Treatment with a combination of 6-thio-dG and V-9302 inhibits
GBM proliferation, an effect that can be detected by hyperpolarized [1-13C]-alanine:
Next,
we examined the effect of treatment with 6-thio-dG, V-9302 or a combination of
6-thio-dG and V-9302 on GBM1 and GBM6 cells. As shown in Fig. 2A-2B, while both
compounds individually inhibited proliferation, the combination of 6-thio-dG
and V-9302 maximally inhibited cell viability. Importantly, the reduction in lactate
production from hyperpolarized [1-13C]-alanine paralleled the effect
of these inhibitors on GBM growth. Representative 13C summed spectra
from GBM6 cells treated with vehicle control (saline), V-9302, 6-thio-dG or the
combination are shown in Fig. 3A. The combination of 6-thio-dG and V-9302
resulted in the highest inhibition of lactate production in both GBM1 and GBM6
models (Fig. 3B).
Hyperpolarized [1-13C]-alanine monitors response
to therapy in vivo: Finally, we performed 2D EPSI studies in rats
bearing orthotopic GBM6 tumors to determine the ability of hyperpolarized [1-13C]-alanine
to assess response to the combination of 6-thio-dG and V-9302. Examination of
metabolic heatmaps (Fig. 4A, 4C) confirmed that the lactate/alanine ratio was
higher in tumor relative to contralateral normal brain, consistent with
previous results in TERT+ oligodendrogliomas4.
Importantly, lactate /alanine heatmaps generated 7 days after treatment with a combination
of 6-thio-dG and V-9302 showed a drop relative to the pre-treatment heatmap (Fig.
4C-4D). There was no change in tumor volume until day 21 (Fig. 4A & 4B; pre-treatment=85mm3,
post-treatment=80mm3; day 21=3mm3), suggesting that lactate
production from hyperpolarized [1-13C]-alanine has the potential to
serve as an early biomarker of GBM response to therapy, prior to anatomical
alterations in vivo.Conclusions
TERT expression drives tumor immortality, making TERT both
a tumor biomarker and a therapeutic target1,2. Our studies
with patient-derived GBM cells and orthotopic tumor xenografts suggest that lactate
production from hyperpolarized [1-13C]-alanine is a non-invasive
metabolic imaging biomarker of TERT expression in GBMs. Combined treatment with
the TERT inhibitor 6-thio-dG and the ASCT2 inhibitor V-9302 abrogates GBM
proliferation, identifying a novel therapeutic opportunity for GBMs.
Importantly, lactate production from hyperpolarized [1-13C]-alanine serves
as a companion imaging biomarker of GBM response to therapy, at early
timepoints prior to volumetric alterations.Acknowledgements
We thank William Byrne for his support in the UCSF
preclinical MR imaging lab. This study was supported by NIH R01CA239288,
Department of Defense W81XWH201055315, UCSF Loglio and NICO initiatives and the
Hana Jabsheh Foundation. The authors acknowledge support from the NIH Center
grant P41EB013598.References
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