Direct electrical coupling between neurons can be the result of both e
lectrotonic current transfer through gap junctions and extracellular f
ields. Intracellular recordings from CA1 pyramidal neurons of rat hipp
ocampal slices showed two different types of small-amplitude coupling
potentials: short-duration (5 ms) biphasic spikelets, which resembled
differentiated action potentials and long-duration (>20 ms) monophasic
potentials. A three-dimensional morphological model of a pyramidal ce
ll was employed to determine the extracellular field produced by a neu
ron and its effect on a nearby neuron resulting from both gap junction
al and electric field coupling. Computations were performed with a nov
el formulation of the boundary element method that employs triangular
elements to discretize the soma and cylindrical elements to discretize
the dendrites. An analytic formula was derived to aid in computations
involving cylindrical elements. Simulation results were compared with
biological recordings of intracellular potentials and spikelets. Fiel
d effects produced waveforms resembling spikelets although of smaller
magnitude than those recorded in vitro. Gap junctional electrotonic co
nnections produced waveforms resembling small-amplitude excitatory pos
tsynaptic potentials. Intracellular electrode measurements were found
inadequate for ascertaining membrane events because of externally appl
ied electric fields. The transmembrane voltage induced by the electric
field was highly spatially dependent in polarity and wave shape, as w
ell as being an order of magnitude larger than activity measured at th
e electrode. Membrane voltages because of electrotonic current injecti
on across gap junctions were essentially constant over the cell and we
re accurately depicted by the electrode. The effects of several parame
ters were investigated: I) decreasing the ratio of intra to extracellu
lar conductivity reduced the field effects; 2) the tree structure had
a major impact on the intracellular potential; 3) placing the gap junc
tion in the dendrites introduced a time delay in the gap junctional me
diated electrotonic potential, as well as deceasing the potential reco
rded by the somatic electrode; and 4) field effects decayed to one-hal
f of their maximum strength at a cell separation of similar to 20 mu m
. Results indicate that the in vitro measured spikelets are unlikely t
o be mediated by gap junctions and that a spikelet produced by the ele
ctric field of a single source cell has the same waveshape as the meas
ured spikelet but with a much smaller amplitude. It is hypothesized th
at spikelets are a manifestation of the simultaneous electric field ef
fects from several local cells whose action potential firing is synchr
onized.