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Impedance Spectroscopy resolves electrical properties into uncorrelatedvariables, as a function of frequency, with exquisite resolution. Separation isrobust and most useful when the system is linear. Impedance spectroscopycombined with appropriate structural knowledge provides insight into pathwaysfor current flow, with more success than other methods. Biological applicationsof impedance spectroscopy are often not useful since so much of biology isstrongly nonlinear in its essential features, and impedance spectroscopy isfundamentally a linear analysis. All cells and tissues have cell membranes andits capacitance is both linear and important to cell function. Measurementsproved straightforward in skeletal muscle, cardiac muscle, and lens of the eye.In skeletal muscle, measurements provided the best estimates of the predominant(cell) membrane system that dominates electrical properties. In cardiac muscle,measurements showed definitively that classical microelectrode voltage clampcould not control the potential of the predominant membranes, that were in thetubular system separated from the extracellular space by substantialdistributed resistance. In the lens of the eye, impedance spectroscopy changedthe basis of all recording and interpretation of electrical measurements andlaid the basis for Rae and Mathias extensive later experimental work. Manytissues are riddled with extracellular space as clefts and tubules, forexample, cardiac muscle, the lens of the eye, most epithelia, and of coursefrog muscle. These tissues are best analyzed with a bidomain theory that arosefrom the work on electrical structure described here. There has been a greatdeal of work since then on the bi-domain and this represents the most importantcontribution to biology of the analysis of electrical structure in my view.

Electrical Structure of Biological Cells and Tissues: impedance spectroscopy, stereology, and singular perturbation theory