Neuroscience Research 36 (2000) 81 – 91
When pyramidal neurons lock, when they respond chaotically, and
when they like to synchronize
R. Stoop
a,
*, K. Schindler
a
, L.A. Bunimovich
b
a
Institut fu ¨r Neuroinformatik, ETHZ/UNIZH, Winterthurerstrasse 190, CH-8057 Zu ¨rich, Switzerland
b
Southeast Applied Analysis Center, Georgia Institute of Technology, Atlanta, GA 30332, USA
Received 20 May 1999; accepted 26 October 1999
Abstract
We give an overview on the locking properties of perturbed regularly firing pyramidal neurons, as a function of perturbation
strength, self-spiking frequency and perturbation frequency. For inhibitory perturbations, instead of locking chaotic response
emerges for a whole range of parameters. This suggests that global synchronization on the set of inhibitory connections may easily
be achieved. © 2000 Elsevier Science Ireland Ltd. All rights reserved.
Keywords: Pyramidal neurons; Regular and chaotic interaction; Global synchronization; Network dynamics
www.elsevier.com/locate/neures
1. Introduction
Suggestions have been made in the past few years
that cortical networks may typically be chaotic (Schiff,
1994). However, at what level of description such a
property can rigorously be established, where it origi-
nates from and what implications for the working brain
this entails are questions that have not been rigorously
answered. As a starting point, we recently were able to
demonstrate numerically that when recurrent excita-
tory – inhibitory connections are activated experimen-
tally, individual neurons exhibit both regular and
chaotic firing patterns (Schindler et al., 1997). In these
in vitro experiments, regularly firing cortical neurons
exhibit chaotic firing patterns when inhibitory pulses
occur with a particular frequency relationship to the
regularly firing neuron. The inhibitory input itself need
not be chaotic at all, indeed it can be as regular as
clockwork and nevertheless produce a chaotic firing
pattern. Rather opposite to the question of chaos, we
have also considered how coherent action/reaction of
the brain could be established, which is believed to be a
prerequisite for higher tasks of the brain. Explaining
the mechanisms of synchronization over large cortical
areas is also of importance for the understanding of
epileptic seizures. Our investigations may contribute to
both groups of questions. We are able to isolate one
possible mesoscopic origin of chaotic behavior in the
brain and we determine the abundance of this behavior
in the natural parameter space of biological pyramidal
neurons. Our findings are that (1) only inhibitory con-
nections can lead to chaotic response; (2) chaotic re-
sponse emerges for a whole range of the parameter
space. As a consequence of the latter property, there is
a tendency towards complete phase-locking among in-
hibitory connections, where formerly independently
chaotically spiking neurons can be brought into stable
spiking patterns bv the application of small couplings.
For a mathematical approach to the underlying general
mechanisms, see Losson and Mackey (1994).
2. Results
In our experiments with pyramidal neurons, slices of
rat neocortex were prepared for in vitro recording.
Following standard techniques, individual pyramidal
neurons in layer 5 of barrel cortex were intracellularly
recorded with sharp electrodes. To induce regular
firing, a constant current was injected into the neurons
(Reyes and Fetz, 1993a,b). The regular firing neuron
* Corresponding author. Tel.: +41-16353063.
E-mail address: ruedi@ini.phys.ethz.ch (R. Stoop)
0168-0102/00/$ - see front matter © 2000 Elsevier Science Ireland Ltd. All rights reserved.
PII:S0168-0102(99)00108-X