Benjamin L Cox1,2,3, Sarah Erickson-Bhatt2,3,4, Joseph M Szulczewski3,4, Kai D Ludwig1, Erin B Adamson1, Robert A Swader2, Suzanne M Ponik4, Kevin W Eliceiri1,2,3,5, and Sean B Fain1,6
1Medical Physics, University of Wisconsin - Madison, Madison, WI, United States, 2Morgridge Institute for Research, Madison, WI, United States, 3Laboratory for Optical and Computational Instrumentation (LOCI), University of Wisconsin - Madison, Madison, WI, United States, 4Cell and Regenerative Biology, University of Wisconsin - Madison, Madison, WI, United States, 5Biomedical Engineering, University of Wisconsin - Madison, Madison, WI, United States, 6Radiology, University of Wisconsin - Madison, Madison, WI, United States
Synopsis
The design and application of a novel bioreactor capable of facilitating
both magnetic resonance spectroscopy (MRS) and optical fluorescence microscopy for
complementary metabolic information is described. Fluorescence lifetime imaging
(FLIM) of nicotinamide adenine dinucleotide (NADH) and hyperpolarized [1-13C]
pyruvic acid (PA) MRS were performed on 3D cell cultures of 4T1 murine breast
cancer cells to study the effects of glucose starvation across cellular and
population scales. The system provides a novel test-bed for simulating cell-matrix
and cell-cell interactions in a 3D microenvironment for investigating
multi-scale cellular metabolism in vitro.
Purpose
Tumor metabolism is an important feature and potential therapy
target in cancer1. However, many aspects of the metabolism of tumor
cells are not fully understood. Fluorescence lifetime imaging (FLIM)2
based optical microscopy and hyperpolarized 13C magnetic resonance
spectroscopy (MRS)3 are methods that enable in vitro and in vivo measurement
of cellular metabolism. FLIM measures the rate of decay of a molecule’s
fluorescence, including the metabolic cofactor nicotinamide adenine
dinucleotide (NADH) whose fluorescence lifetime depends on its protein binding
state allowing visualization of metabolic processes that are dominant within a
cell4. Hyperpolarized MRS most commonly uses hyperpolarized [1-13C]
pyruvic acid to measure metabolic products of glycolysis, especially lactate
production. As these techniques occur on very different scales they can yield
complementary information at the individual cell and entire population scales (Figure 1). Here, we present a
multi-modal and multi-scale study of the effects of glucose starvation on 4T1 murine
breast cancer cell metabolism using a novel MRI and optical microscopy
compatible bioreactor capable of combined MRS and NADH FLIM in 3D cell cultures comprised of collagen gels that mimic the
tumor microenvironment.Methods
The bioreactor (Figure
2) was designed using SolidWorks (Dassault Systemes, Velizy-Villacoublay,
France). The experimental setups for both optical microscopy and MRS experiments are shown in Figure 3. Temperature
feedback is achieved with two independent sample probes for optical imaging and
with an MR-compatible temperature probe for MRS experiments.
4T1 murine breast cancer cells5 in normal growth media
were compared to 4T1 cells in glucose-free media to study the effects of
glucose starvation. Collagen gels were polymerized as previously described6.
In brief, collagen was neutralized with 2x HEPES buffer, then further
diluted to 2mg/mL with cells and media. Approximately 8 million cells were used
per replicate. After polymerization of collagen, cells were allowed to incubate
for 1-2 hours prior to experiments. For the glucose-free state, glucose-free
media was used for this incubation period. Optical imaging was performed with a
20x air objective on a custom multiphoton laser scanning system built around an
inverted Nikon TE300 microscope7. MRS experiments were performed in
a 4.7T preclinical MRI scanner (Agilent, Palo Alto, CA).
For MRS experiments, 30µL aliquots of [1- 13C]PA
(Cambridge Isotope Laboratories Inc., Tewksbury, MA) and 15mM trityl radical
(Ox063, GE Healthcare) were polarized in a Hypersense polarizer (Oxford
Instruments) for ~1 hour8. Samples were dissoluted with 4mL of
solvent containing 1.2mL 426mM NaOH, 1.4mL 400 mM Tris buffer and 1.4mL 250mg/L
EDTA. [1-13C]PA was drawn off and 400µL were rapidly injected into
the sample volume of the bioreactor. Dynamic global spectra (FA=10º, TR=3000ms) were acquired
with a dual-tuned 1H/13C volume coil (Doty Scientific,
Columbia, SC). N=9 FLIM (3 each from 3 bioreactor preparations) and N=5 MRS (2
bioreactors were prepared for each state in addition to the 3 used for combined
studies) measurements were aquired for each state.
Results
MRS results showed a significant decrease (p<0.01) in
lactate production of 4T1 cells in the glucose-free state compared to the normal
state (Figure 4). NADH lifetime
results showed a corresponding increase in mean NADH lifetime of 4T1 cells from
the normal state to the glucose-free state (p<0.01) (Figure 4). 4T1 cells in the glucose-free state are also less
densely clumped together and have a rounder overall shape than those in the
normal state (Figure 4B vs. 4D).Discussion
Here, multi-scale imaging was used to investigate the
juxtaposition of cellular and population level changes in metabolism in
response to glucose starvation. Changes in lactate production at the population
level, coincided with changes in the chemical bound state of NADH and
morphological changes at the cellular level between normal and glucose-starved
4T1 cells. Both changes suggest a shift away from glycolysis in the
glucose-free state. This could indicate other metabolic processes at work,
cells beginning to die of starvation or both, as the 4T1 strain is known to be
a metastatic cell line that heavily depends on glycolysis9. Future
work will aim to investigate more precise inhibition of metabolic pathways in
concert with markers of apotosis and recovery after reintroduction of glucose
and lactate substrates in a wider range of breast cancer cell lines. Conclusion
Hyperpolarized [1-13C] PA MRS (global metabolism)
and NADH fluorescence lifetime (cellular metabolism) were performed on 4T1 murine
breast cancer cells under normal and glucose-starved conditions, showing a
shift away from glycolysis in the glucose-starved state at both the population
and cellular scales. This system represents a novel approach to achieving
multimodal and multiscale measurement of metabolism in 3D cell cultures.Acknowledgements
Funding for this project was provided by The Morgridge
Institute for Research, the Departments of Medical Physics and Radiology at
UW-Madison, UL1TR000427 to UW ICTR from NIH/NCATS and the AAPM 2014
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