Double diffusion encoding MRS (DDE-MRS) was shown to be highly sensitive to cell microstructure, in particular cell fiber diameter. Here we revisit the ability of DDE-MRS to probe microstructure by quantifying DDE-MRS signal for six metabolites with high accuracy. We show that signal modulation differs for neuronal and glial metabolites, yielding larger fiber diameters for glia when using a model of isotropically oriented cylinders, as previously reported. However, fiber diameters appear overestimated. Further data acquisition and modeling suggests DDE-MRS is not only sensitive to fiber diameter but also to other microstructural features, such as cell body diameter or fiber length.
Quality of DDE-MRS spectra: Amplitude modulation of spectra can be observed in Figure 1-A. NAA, Ins, taurine (Tau), total creatine (tCr), total choline (tCho), lactate (Lac) and MM were reliably quantified (CRLBs<5%) using LCModel [10] as shown in the spectral decomposition displayed in Figure 1-B. Note the very low s.d. that could be achieved in our experiments.
Sensitivity to cellular compartments: Figure 2 shows the fitting of experimental data for all metabolites while A and B coefficients are reported in Table 1. It can be seen that the amplitude modulation of the DDE-MRS signal -reflected here by B- is larger for NAA (neuronal metabolite) than for non-specific (tCr and Tau) or glial markers (tCho and Ins). This is consistent with the fact that neurons are expected to exhibit narrower fibers / larger CSA than glia [5, 11]. Interestingly, lactate exhibits stronger signal attenuation, which could be explained by the intrinsically fast diffusion of lactate (which is a small metabolite), but also by the contribution of a significant extracellular lactate pool with a rather "free-like" diffusion behavior [12]. In line with this idea, lactate exhibits less pronounced angular amplitude modulation than intracellular metabolites.
Estimating microstructure: Heatmaps based on least square residuals for each metabolite as compared to DDE-MRS simulations, as a function of Dfree and d, are presented in Figure 3. They show similar aspect for all metabolites except lactate. Diameters corresponding to the best fits are in the 3-5 µm range (Table 1), which is unrealistically large (for example, astrocytic processes are ~1-µm in diameter) [13], and Dfree is unusually low (~0.2 µm²/ms, while they are generally estimated to be 0.3-0.4 µm²/ms). One possible origin might be the contribution of cell bodies: for example, assuming that 20% of the signal originates from 10-µm diameter spherical cell bodies and 80% for 1-µm diameter fibers results in some angular dependency that is relatively close to experimental data (Figure 4-A). Another possible origin might be the long-range fiber structure (e.g. branching and finite length), as shown in Fig. 4-B presenting simulations in 1-µm diameter and 30-µm length cylinders (without spheres). Very interestingly, both configurations predict higher signal intensity at 180° than at 0/360°, especially when decreasing TM. We performed a new series of experiments at shorter TM, which appear to experimentally confirm this trend (Figure 4-C). This emphasizes the need to go beyond a simple picture of infinite-length cylinders.
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