Journal of ComputationalNeuroscience3, 199-223 (1996)
© 1996 KluwerAcademicPublishers. Manufacturedin The Netherlands.
Dissection and Reduction of a Modeled Bursting Neuron
R.J. BUTERA, JR. AND J.W. CLARK, JR.
Dept. of Electrical and Computer Engineering, MS 366, Rice University, Houston, TX 77251-1892
rbutera@ece.fice.edu
J.H. BYRNE
Dept. of Neurobiology and Anatom.3; University of Texas Medical School, Houston, TX 77030
jbyrne@nbal9.med.uth.tmc.edu
Received September 5, 1995; Revised February 1, 1996; Accepted February 13, 1996
Action Editor: John Rinzel
Abstract. An 11-variable Hodgkin-Huxley type model of a bursting neuron was investigated using numerical
bifurcation analysis and computer simulations. The results were applied to develop a reduced model of the underlying
subthreshold oscillations (slow-wave) in membrane potential. Two different low-order models were developed: one
3-variable model, which mimicked the slow-wave of the full model in the absence of action potentials and a second
4-variable model, which included expressions accounting for the perturbational effects of action potentials on the
slow-wave. The 4-variable model predicted more accurately the activity mode (bursting, beating, or silence) in
response to application of extrinsic stimulus current or modulatory agents. The 4-variable model also possessed
a phase-response curve that was very similar to that of the original 11-variable model. The results suggest that
low-order models of bursting cells that do not consider the effects of action potentials may erroneously predict
modes of activity and transient responses of the full model on which the reductions are based. These results also
show that it is possible to develop low-order models that retain many of the characteristics of the activity of the
higher-order system.
Keywords: nonlinear dynamics, bifurcation, bursting, model reduction
1. Introduction
The endogenous oscillations in the membrane po-
tential of excitable membranes have been the sub-
ject of extensive analyses using mathematical mod-
eling and computer simulation approaches. Among
the most detailed models are those of cells whose os-
cillations in membrane potential exhibit bursting ac-
tivity: a slow oscillation in membrane potential that
alters between a silent state and a bursting state. The
bursting state is characterized by the firing of a suc-
cession of action potentials. The most extensively
modeled bursting cells are pancreatic fl-cells (Chay
and Keizer, 1983; Chay, 1990; Keizer and Man-
gus, 1989; Sherman et al., 1988) and neuron R15 in
Aplysia (Bertram, 1993; Butera et al., 1995; Canavier
et al., 1991; Plant and Kim, 1976). Electrophysi-
ological investigations over the past several decades
have revealed a rich repertoire of biophysical mech-
anisms. Recent models simulate not only endoge-
nous activity and changes in behavior produced by
extrinsic currents, but also simulate changes in en-
dogenous activity as the concentration of extrinsic
modulatory agents are varied (Bertram, 1993; Butera
et al., 1995; Keizer and Mangus, 1989). To achieve
this level of accuracy and predictability the models