Interpretation of time constant and electrotonic length estimates in multicylinder or branched neuronal structures.
1. We have investigated the theoretical and practical problems associated with the interpretation of time constants and the estimation of electrotonic length with equivalent cylinder formulas for neurons best represented as multiple cylinders or branched structures. Two analytic methods were used to compute the time constants and coefficients of passive voltage transients (and time constants of current transients under voltage clamp). One method, suitable for simple geometries, involves analytic solutions to boundary value problems. The other, suitable for neurons of any geometric complexity, is an algebraic approach based on compartmental models. Neither of these methods requires the simulation of transients. 2. We computed the time constants and coefficients of voltage transients for several hypothetical neurons and also for a spinal motoneuron whose morphology was characterized from serial reconstructions. These time constants and coefficients were used to generate voltage transients. Then exponential peeling, nonlinear regression, and transform methods were applied to these transients to test how well these procedures estimate the underlying time constants and coefficients. 3. For a serially reconstructed motoneuron with 732 compartments, we found that the theoretical and peeled tau 0 values were nearly equal, but the theoretical tau 1 was much larger than the peeled tau 1. The theoretical tau 1 could not be peeled because it was associated with a coefficient, C1, that had a very small value. In fact, there were 156 time constants between 1.0 and 6.0 ms, most of which had very small coefficients; none had a coefficient larger than 2% of the signal. The peeled value of tau 1 (called tau 1 peel) can be viewed as some sort of a weighted average of the time constants having the largest coefficients. 4. We studied simple hypothetical neurons to determine what interpretation could be applied to the multitude of theoretical time constants. We found that after tau 0, there was a group of time constants associated with eigenfunctions that were odd (or approximately odd) functions with respect to the soma. These time constants could be interpreted as "equalizing" time constants along particular paths between different pairs of dendritic terminals in the neuron. After this group of time constants, there was one that we call tau even because it was associated with an eigen-function that was approximately even with respect to the soma. This tau even could be interpreted as an equalizing time constant for charge equalization between proximal membrane (soma and proximal dendrites) and distal membrane (including all distal dendrites).4=
Holmes
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Mathematical Research Branch, National Institute of Diabetes and Digestive and Kidney Diseases, National Institutes of Health, Bethesda, Maryland 20892.
Segev
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Rall
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D000818:Animals; D003712:Dendrites / Q000502:physiology* / Q000648:ultrastructure; D004594:Electrophysiology; D008433:Mathematics; D008959:Models, Neurological; D008961:Models, Structural; D009474:Neurons / Q000166:cytology* / Q000502:physiology* / Q000648:ultrastructure; D013997:Time Factors