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* University of Kentucky, College of Medicine, Lexington, Kentucky; University of Maryland, School of Nursing, Baltimore, Maryland; University of Auckland, Auckland, New Zealand; Rochester Institute of Technology, Rochester, New York; and ¶ University of Maryland, Baltimore County, Maryland
Correspondence: Address reprint requests to Leighton T. Izu, PhD, Institute of Molecular Medicine, University of Kentucky BBSRB, Rm. B257, 741 South Limestone St., Lexington, KY 40536-0509. Tel.: 859-323-6882; E-mail: leightonizu{at}uky.edu; web site: http://www.mc.uky.edu/imm/; or Ye Chen-Izu, PhD, Institute of Molecular Medicine, University of Kentucky BBSRB, Rm. B257, 741 South Limestone St., Lexington, KY 40536-0509, Tel.: 859-323-6882; E-mail: YeChen-Izu{at}uky.edu; web site: http://www.mc.uky.edu/imm/.
The clustering of ryanodine receptors (RyR2) into functional Ca2+ release units is central to current models for cardiac excitation-contraction (E-C) coupling. Using immunolabeling and confocal microscopy, we have analyzed the distribution of RyR2 clusters in rat and ventricular atrial myocytes. The resolution of the three-dimensional structure was improved by a novel transverse sectioning method as well as digital deconvolution. In contrast to earlier reports, the mean RyR2 cluster transverse spacing was measured 1.05 µm in ventricular myocytes and estimated 0.97 µm in atrial myocytes. Intercalated RyR2 clusters were found interspersed between the Z-disks on the cell periphery but absent in the interior, forming double rows flanking the local Z-disks on the surface. The longitudinal spacing between the adjacent rows of RyR2 clusters on the Z-disks was measured to have a mean value of 1.87 µm in ventricular and 1.69 µm in atrial myocytes. The measured RyR2 cluster distribution is compatible with models of Ca2+ wave generation. The size of the typical RyR2 cluster was close to 250 nm, and this suggests that 
100 RyR2s might be present in a cluster. The importance of cluster size and three-dimensional spacing for current E-C coupling models is discussed.
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