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J Neurophysiol 83: 3310-3322, 2000;
0022-3077/00 $5.00
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The Journal of Neurophysiology Vol. 83 No. 6 June 2000, pp. 3310-3322
Copyright ©2000 by the American Physiological Society

Linear to Supralinear Summation of AMPA-Mediated EPSPs in Neocortical Pyramidal Neurons

Jilda S. Nettleton1,3 and William J. Spain1,2,3

 1Veterans Affairs Puget Sound Health Care System, Seattle 98108 and  2Department of Neurology and  3Department of Physiology and Biophysics, University of Washington School of Medicine, Seattle, Washington 98195

Nettleton, Jilda S. and William J. Spain. Linear to Supralinear Summation of AMPA-Mediated EPSPs in Neocortical Pyramidal Neurons. J. Neurophysiol. 83: 3310-3322, 2000. It has been hypothesized that voltage-sensitive conductances present on the dendrites of neurons can influence summation of excitatory postsynaptic potentials (EPSPs) and hence affect how neurons compile information. Greater than linear summation of EPSPs has been postulated to facilitate coincidence detection by cortical neurons. This study examined whether the summation of subthreshold AMPA-mediated EPSPs generated on layer V neocortical pyramidal neurons in vitro was linear and if any nonlinearities could be attributed to dendritic conductances. Evoked EPSPs (1-12 mV) were recorded somatically by means of intracellular sharp electrodes in the presence of 100 µM amino-5-phosphonopentanoic acid (AP-5) and 3 µM bicuculline. Two independent EPSPs were evoked by a stimulating electrode in layer I and another in layers III-V. The areas of stimulation were isolated from each other by a horizontal cut below layer I. By subtracting the algebraic sum of the individual EPSPs from the evoked response when both EPSPs were evoked simultaneously, we determined that they summed linearly to supralinearly. Supralinear summation was more likely when the soma was hyperpolarized by DC current injection. Summation was predominantly linear when postsynaptic conductances (i.e., Na+ and Ca2+) were blocked with intracellular QX-314. The supralinear summation of EPSPs (without QX-314) decreased as the time between inputs was increased from 0 to 30 ms. To determine the role of dendrites in nonlinear summation, we substituted a current pulse (simulated EPSP) delivered at the soma for either or both of the evoked EPSPs. Simulated EPSPs combined with either an evoked EPSP or another simulated EPSP showed significantly less supralinear summation than two evoked EPSPs, indicating that the dendritic conductances were largely responsible for the observed supralinear summation.




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