10× EExM visualization of variable phosphorylation
of RyRs
during pathology and acute stimulation. (A) Overlay of 10× EExM
images of RyR (gray) and pS2808 (purple) at the surface of a rat ventricular
cardiac myocyte. Inset shows the equivalent view in a region ∼5
μm below the cell surface. (B) Magnified view of nanodomains
near (i) the cell surface and (ii) cell interior visually illustrates
that punctate pS2808 (purple) labeling densities, observed in fewer
numbers, follow either close overlap or proximity with RyR puncta
(gray). (C) The centroids of the RyR (gray) and pS2808 puncta (purple)
were localized and paired with the nearest RyR within a 30 nm 3D distance
(dashed line) to identify RyRs which were phosphorylated at the Ser2808
site. Raw data of region illustrated in inset. An example of unpaired
pS2808 marker is shown with an arrowhead. Magnified views of the overlays
of subsurface nanodomains in cells from the right ventricles of (D)
control and (E) MCT rats show a minor fraction of RyRs which associate
pS2808 labeling. The same comparison in the cell interior regions
illustrates a similar fraction of RyRs with Ser2808 phosphorylation
in (F) control nanodomains, while a greater proportion of RyRs, despite
smaller cluster size, coincide with pS2808 labeling in (G) MCT-RV
nanodomains. (H) Percentage histograms of the mean fraction of RyRs
determined to be phosphorylated at Ser2808 (Pphos) in the subsurface nanodomains containing ≥4 RyRs
illustrate near-identical distributions (mean ± SD of Pphos of 0.31 ± 0.10 and 0.30 ± 0.12; p > 0.05; df = 20 cells; Bonferroni-corrected Mann–Whitney t test) for both control (purple) and MCT-RV (blue). (I)
Cell interior nanodomains in MCT-RV (blue; 0.45 ± 0.07) showed
a rightward shifted Pphos distribution
compared to control (purple; 0.27 ± 0.09 ; p = 0.022, df = 19). Pphos distributions
for both subsurface and interior nanodomains appeared to show a right-shift
in control cells stimulated with isoproterenol and simultaneous electrical
pacing at 1 Hz (insets). (J) Histograms of the 2D pS2808 localization
density as a function of the distance within the subsurface nanodomain
relative to its boundary in control (purple line), stimulated (black),
and MCT-RV (blue) cells. (K-i) In control cells, RyRs phosphorylated
at Ser2808 are located, on average, uniformly throughout the nanodomain.
(K-ii) Stimulation with isoproterenol and electrical pacing appeared
to approximately double the pS2808 density uniformly across the nanodomain.
(K-iii) In MCT-RV, a gradient of pS2808 density is seen from the edge
of the nanodomain inward, allowing a subdomain of potentiated RyRs
to be maintained at the center of the nanodomain (circle). Scale bars:
(A) 200 nm, (B–G) 50 nm.