Abstract
One way to achieve amplification of distal synaptic inputs on a dendritic tree is to scale the amplitude and/or duration of the synaptic conductance with its distance from the soma. This is an example of what is often referred to as “dendritic democracy”. Although well studied experimentally, to date this phenomenon has not been thoroughly explored from a mathematical perspective. In this paper we adopt a passive model of a dendritic tree with distributed excitatory synaptic conductances and analyze a number of key measures of democracy. In particular, via moment methods we derive laws for the transport, from synapse to soma, of strength, characteristic time, and dispersion. These laws lead immediately to synaptic scalings that overcome attenuation with distance. We follow this with a Neumann approximation of Green’s representation that readily produces the synaptic scaling that democratizes the peak somatic voltage response. Results are obtained for both idealized geometries and for the more realistic geometry of a rat CA1 pyramidal cell. For each measure of democratization we produce and contrast the synaptic scaling associated with treating the synapse as either a conductance change or a current injection. We find that our respective scalings agree up to a critical distance from the soma and we reveal how this critical distance decreases with decreasing branch radius.


















Similar content being viewed by others
References
Abbott, L. F., Fahri, E., & Gutmann, S. (1991). The path integral for dendritic trees. Biological Cybernetics, 66, 49–60.
Agmon-Snir, H. (1995). A novel theoretical approach to the analysis of dendritic transients. Biophysical Journal, 69(5), 1633–1656.
Andersen, P., Silfvenius, H., Sundberg, S. H., & Sveen, O. (1980) A comparison of distal and proximal dendritic synapses on CA1 pyramids in guinea-pig hippocampal slices in vitro. Journal of Physiology, 307, 273–299.
Andrasfalvy, B. K., & Magee, J. C. (2001). Distance-dependent increase in AMPA receptor number in the dendrites of adult hippocampal CA1 pyramidal neurons. Journal of Neuroscience, 21, 9151–9159.
Bressloff, P., & Coombes, S. (1997). Physics of the extended neuron. International Journal of Modern Physics B, 11, 2343–2392.
Burke, W. (1957). Spontaneous potentials in slow muscle fibres of the frog. Journal of Physiology, 135(3), 511–21.
Carnevale, N. T., & Hines, M. L. (2006). The NEURON Book. Cambridge: Cambridge University Press.
Coombes, S., Timofeeva, Y., Svensson, C.-M., Lord, G. J., Josić, K., Cox, S. J., et al. (2007). Branching dendrites with resonant membrane: a “sum-over-trips” approach. Biological Cybernetics, 97, 137–149.
Cuntz, H., Borst, A., & Segev, I. (2007). Optimization principles of dendritic structure. Theoretical Biology and Medical Modelling, 4, 21.
De Schutter, E., & Bower, J. M. (1994). Simulated responses of cerebellar Purkinje cells are independent of the dendritic location of granule cell synaptic inputs. Proceedings of the National Academy of Sciences of the United States of America, 91, 4736–4740.
Fatt, P., & Katz, B. (1951). An analysis of the end-plate potential recorded with an intracellular electrode. Journal of Physiology, 115(3), 320–70.
Golding, N., Mickus, T., Katz, Y., Kath, W., & Spruston, N. (2005). Factors mediating powerful voltage attenuation along CA1 pyramidal neuron dendrites. The Journal of Physiology, 568(1), 69–82.
Häusser, M. (2001). Synaptic function: Dendritic democracy. Current Biology, 11, R10–R12.
Iansek, R., & Redman, S. J. (1973). The amplitude, time course and charge of unitary excitatory post-synaptic potentials evoked in spinal motoneurone dendrites. Journal of Physiology, 234, 665–688.
Jack, J., Noble, D., & Tsien, R. (1975). Electric current flow in excitable cells. Oxford: Oxford University Press.
Jack, J. J., Redman, S. J., & Wong, K. (1981). The components of synaptic potentials evoked in cat spinal motoneurones by impulses in single group Ia afferents. Journal of Physiology, 321, 65–96.
Koch, C. (1999). Biophysics of computation: Information processing in single neurons. Oxford: Oxford University Press.
London, M., & Segev, I. (2001). Synaptic scaling in vitro and in vivo. Nature Neuroscience, 4, 853–854.
Magee, J., & Cook, E. (2000). Somatic EPSP amplitude is independent of synapse location in hippocampal pyramidal neurons. Nature Neuroscience, 3, 895–903.
Magee, J. C. (2000). Dendritic integration of excitatory synaptic input. Nature Reviews Neuroscience, 1, 181–190.
Migliore, M., Ferrante, M., & Ascoli, G. A. (2005). Signal propagation in oblique dendrites of CA1 pyramidal cells. Journal of Neurophysiology, 94, 4145–4155.
Nicholson, D., Trana, R., Katz, Y., Kath, W., Spruston, N., & Geinisman, Y. (2006a). Distance-dependent differences in synapse number and AMPA receptor expression in hippocampal CA1 pyramidal neurons. Neuron, 50(3), 431–442.
Nicholson, D. A., Trana, R., Katz, Y., Kath, W. L., Spruston, N., & Geinisman, Y. (2006b). Distance-dependent differences in synapse number and AMPA receptor expression in hippocampal CA1 pyramidal neurons. Neuron, 50(3), 431–442.
Rall, W. (1964). Theoretical significance of dendritic trees for neuronal input–output relations. In R. F. Reiss (Ed.), Neural theory and modeling. Stanford: Stanford University Press.
Rall, W., & Agmon-Snir, H. (1998). Cable theory for dendritic neurons. In C. Koch & I. Segev (Eds.), Methods in Neuronal Modeling (2nd ed., pp. 27–92). Cambridge, MA: MIT Press.
Rumsey, C. C., & Abbott, L. F. (2004). Equalization of synaptic efficacy by activity- and timing-dependent synaptic plasticity. Journal of Neurophysiology, 91, 2273–2280.
Rumsey, C. C., & Abbott, L. F. (2006). Synaptic democracy in active dendrites. Journal of Neurophysiology, 96(5), 2307–2318.
Timofeeva, Y., Lord, G. J., & Coombes, S. (2006a). Dendritic cable with active spines: A modeling study in the spike-diffuse spike framework. Neurocomputing, 69, 1058–1061.
Timofeeva, Y., Lord, G. J., & Coombes, S. (2006b). Spatio-temporal filtering properties of a dendritic cable with active spines. Journal of Computational Neuroscience, 21, 293–306.
Tuckwell, H. C. (1988a). Introduction to theoretical neuro biology: Linear cable theory and dendritic structure, vol. I of Cambridge studies in mathematical biology. Cambridge: Cambridge University Press.
Tuckwell, H. C. (1988b). Introduction to theoretical neurobiology: Nonlinear and stochastic theories, vol. II of Cambridge studies in mathematical biology. Cambridge: Cambridge University Press.
Acknowledgements
The work in this paper is supported through Engineering and Physical Sciences Research Council (EPSRC) Grant No. GR/S60914/01. S Coombes would also like to acknowledge ongoing support from the EPSRC through the award of an Advanced Research Fellowship, Grant No. GR/R76219. K Josić was supported by NSF grant DMS-0604429, and an Advanced Research Program/Advanced Technology Program grant.
Author information
Authors and Affiliations
Corresponding author
Additional information
Action Editor: David Terman
Rights and permissions
About this article
Cite this article
Timofeeva, Y., Cox, S.J., Coombes, S. et al. Democratization in a passive dendritic tree: an analytical investigation. J Comput Neurosci 25, 228–244 (2008). https://doi.org/10.1007/s10827-008-0075-9
Received:
Revised:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s10827-008-0075-9