The initiation site in neurons is where the excitatory and inhibitory
inputs sum to generate action potentials. It is generally considered t
o be at a fixed location, typically at the axon hillock or initial seg
ment, although action potentials, or impulses, could in theory arise a
t a site that shifts dynamically. The data reported here show that the
initiation site can shift in a graded manner, by as much as 175 mu m,
depending on the level of neuronal excitation. Laser axotomy reveals
that the Anterior Pagoda (AP) neuron of the leech is excitable within
the synaptic neuropil before its axon bifurcates. Using an electrophys
iological technique to measure relative delays in impulses arriving at
different sites, we have found that depolarization, either by applied
current or by synaptic input, can shift the site of impulse initiatio
n in the cell proximally toward the soma and neurites receiving synapt
ic input. Impulse initiation in this region should enhance the efficac
y of inputs synapsing there. Conversely, hyperpolarization can shift t
he initiation site distally. A shifting initiation site, therefore, ma
y be a mechanism by which synaptic inputs can rapidly enhance or suppr
ess the active response of the AP neuron to other synaptic inputs.