
L AB V ISIT 8
☞ L AB V ISIT 8
Steady States and Periodicity
in a Squid Neuron
S
TEADY STATES and periodic orbits are the two simplest forms of behavior
for discrete and continuous dynamical systems. For two-dimensional systems of
differential equations, all bounded solutions must converge either to periodic
orbits or to sets containing equilibrium points and perhaps connecting arcs,
according to the Poincar
´
e-Bendixson Theorem. In this Lab Visit, we will see
evidence of coexisting attractors in a biological system, one a steady state and the
other a periodic orbit.
Periodic orbits are ubiquitous in biological systems, and interruption of pe-
riodicity is often a symptom of malfunction or disease. In physiology alone, a
significant proportion of medical problems exhibit abnormal dynamical behav-
ior, including tremor and Parkinsonism, respiratory and cardiac disorders, sleep
apnea, epileptic seizures, migraine, manic-depressive illness, endocrinological ir-
regularities, and hiccups. (For the last, see (Whitelaw et al., 1995).)
In particular, the nervous systems of animals thread together a vast hierarchy
of oscillatory processes. Cyclic behavior goes on at many scales, and neurophysi-
ologists search for mechanisms that form the control point of these processes. The
eventual goal is to find the critical parameters that affect the temporal pattern of
a given process, and to determine how to recover from an abnormal setting.
The experiment reported here demonstrated the possibility of intervening
in a biological system to move a trajectory from one of the coexisting attractors
to the other. The system is a neuronal circuit that is well-studied in the scientific
literature, the giant axon from a squid. The axon is the part of the neuron that is
responsible for transmitting information in the form of pulses to other neurons.
The squid giant axon is often chosen because of its size, and the fact that it can
be made to continue firing in a fairly natural manner after it is dissected from the
squid.
The axon was bathed in an artificial seawater solution with abnormally
low calcium concentration, held at approximately 22
◦
C. Periodic firing was
Guttman, R., Lewis, S., Rinzel, J., 1980. Control of repetitive firing in squid axon
membrane as a model for a neuroneoscillator. J. Physiology 305:377–395.
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