Dynamics and Complexity of FitzHugh-Nagumo Neuronal Systems

Mada Sanjaya W. S
1
, Muhammad Yusuf
1
, Agus Kartono
1
, Irzaman
2


1
Theoretical Physics Division, Department of Physics, Faculty of Mathematical and Natural
Sciences, Institut Pertanian Bogor, Bogor 16680, Indonesia.

2
Applied Physics Division, Department of Physics, Faculty of Mathematical and Natural
Sciences, Institut Pertanian Bogor, Bogor 16680, Indonesia.
e-mail : madasws@gmail.com

Abstract

System of signals propagation from one neuron to another represent event of very
complex electrochemical mechanism This work adresses the dynamics and
complexity of neuron mathematical models. The aim is first the understanding of the
biological meaning of existing mathematical systems concerning neurons such as
Hodgkin-Huxley or FitzHugh-Nagumo models. The local stability and the numerical
asymptotic analysis of FitzHugh-Nagumo model are then developed in order to
comprehend the dynamic evolution of a single FitzHugh-Nagumo neuron. This
examination naturally comes to the study of neuron networks. The analysis of these
networks uses the synchronization theory via connections between neurons and can
give rise to emergent properties and self-organization. Our result leads to a classical
law which describes many self-organized complex systems like earthquakes,
linguistic or urban systems. This has been performed using numerical tools.

Keyword: Hodgkin-Huxley models, FitzHugh-Nagumo, action potentials,
synchronization, chaos.

Introduction

Understanding the mechanisms of the
propagation of the nerve activity is one of
the fundamental problems in biophysics.
The first detailed quantitative
measurements of the ionic currents were
performed by Hodgkin and Huxley in the
early 50-s (Hodgkin & Huxley, 1952).
Using the voltage clamp technique, they
were able to measure the kinetics of Na+
and K+ currents in the squid giant axon.
This led them to a set of differential
equations which describe the dynamics of
the action potential. Furthermore, by
combining these equations with the cable
equation for spreading of current in the
axon they were able to calculate the shape
and velocity of the propagating action
potentials (Hodgkin & Huxley, 1952). The
predictions of their model turned out to be
in a remarkably good agreement with the
experimental observations.
Hodgkin-Huxley model for the action
potential of a space clamped squid axon is
de_ned by the four dimensional vector
field. with variables (v; m; n; h) that
represent membrane potential, activation
of a sodium current, activation of a
potassium current, and inactivation of the
sodium current and a parameter I that
represents injected current into the space-
clamped axon. Although there are
improved models the Hodgkin-Huxley
model remains the paradigm for
conductance-based models of neural
system. FitzHugh was the first
investigator to apply qualitative phase-
plane methods to understanding the
Hodgkin-Huxley model. To make





2
headway in gaining analytic insight,
FitzHugh first considered the variables
that change most rapidly, viewing all
others as slowly varying parameters of the
system. In this way he derived a reduced
two-dimensional system that could be
viewed as a phase plane. From the
Hodgkin-Huxley equations FitzHugh
noticed that the variables V and m change
more rapidly than h and n, at least during
certain time intervals. By arbitrarily
setting h and n to be constant we can
isolate a set of two equations which
describe a two-dimensional (V; m) phase
plane. The elegance of applying phase
plane methods and reduced systems of
equations to this rather complicated
problem should not be underestimated.
Determining the dynamical behavior
of an ensemble of coupled neurons is an
important problem in computational
neuroscience. Commonly used models for
the study of individual neuron which
display spiking behavior (FitzHugh, 1961;
Nagumo, 1962; Hodgkin, 1952). From the
very beginning of the research in the field
of computational neuroscience, people
deal with single neuron and its behavior.
Present trends of research include
investigation of the behavior of neurons
considered in a network and their way to
fire synchronously. It is assumed that the
activities in the brain are synchronous and
underlying interests for synchronization of
nonlinear oscillators in physical and
biological systems range from novel
communication strategies to understand
how large and small neural assemblies
efficiently and sensitively achieve desired
functional goal. In recent years, there has
been tremendous interest for the study of
the synchronization of chaotic systems.
The phenomenon of synchronism gives
rise to different dynamical behaviors such
as chaotic synchronization etc (Belykh, I
& Shilnikov, A., 2008). In (Mishra, 2006),
nonlinear dynamical analysis on single
and coupled modified FitzHugh-Nagumo
model under steady current stimulation is
carried out. Also the effect of parameter
variation on its behavior is investigated.

Mathematical Models

Basics Models of Neuron

To describe the evolution of the
membrane potential V in the squid giant
axon (Hodgkin, A.L & Huxley,A.F.,
1952) developed the following
conductance-based model:

| |
eks L L k K Na Na
I v v g v v v n g v v h m v g
C dt
dv
+ ÷ + ÷ + ÷ ÷ = ) ( ) )( ( ) ( ) (
1
4 3

n v n v
dt
dn
n n
) ( ) 1 )( (   ÷ ÷ =
m v m v
dt
dm
m m
) ( ) 1 )( (   ÷ ÷ = (1)
h v h v
dt
dh
h h
) ( ) 1 )( (   ÷ ÷ =

The peak conductances g, reversal
potentials E and rate coefficients and of
the individual
currents and gating variables m, n and h
are given by :
1 10 / ) 40 (
) 1 )( 40 ( 1 . 0 ) (
÷ +
÷ + =
v
m
e v v  ,
18 / ) 65 (
4 ) (
+ ÷
=
v
m
e v 
20 / ) 65 (
07 . 0 ) (
+ ÷
=
v
h
e v  ,
1 10 / ) 35 (
) 1 ( ) (
÷ + ÷
+ =
v
h
e v 
1 10 / ) 55 (
) 1 )( 55 ( 01 . 0 ) (
÷ + ÷
÷ + =
v
n
e v v  ,
80 / ) 65 (
125 . 0 ) (
+ ÷
=
v
n
e v 
For I = 0, the membrane potential settles
into a resting potential of about V = −65
mV.





3
And with the g
Na
= 120, g
K
= 36, g
L
= 0.3
mmho cm
-2
, v
Na
= 50 mV, v
K
= -77 mV
and v
L
= -54.4 mV.
















(a) (b)
Figure 1. Dinamical Systems in Hodgkin_huxley models at I = 0 ; (a) Action potentials at
t = 5 ms, (b) variable m, h and n at t = 5 ms.

FitzHugh-Nagumo Models of Single
Neuron

In 1961 FitzHugh proposed to
demonstrate that the Hodgkin-Huxley
model belongs to a more general class of
systems that exhibit the properties of
excitability and oscillations. As a
fundamental prototype, the van der Pol
oscillator was an example of this class,
and FitzHugh therefore used it (after
suitable modification). A similar approach
was developed independently by Nagumo
in 1962. FitzHugh proposed the following
equations:

) (
3
1
3
bw a v
dt
dw
I w v v
dt
dv
÷ + =
+ ÷ ÷ =

(2)

In these equations the variable v
represents the excitability of the system
and could be identified with voltage
(membrane potential in the axon); w is a
recovery variable, representing combined
forces that tend to return the state of the
axonal membrane to rest. Finally I is the
applied stimulus that leads to excitation
(such as input current), or rectangular
pulses.

Local Dynamics and Bifurcation

Let us consider the FitzHugh-Nagumo
system (2), Equilibria are given by the
following system :

0 / , 0 / = = dt dw dt dv (3)

The nature of this equilibria is
given by the study of the eigenvalues of
the jacobian matrix J of this system :

(
¸
(

¸

÷
÷ ÷
=
  b
v
J
1 1
2
(4)

We obtain the following polynomial
equation :

0 ) ˆ ( ) ˆ 1 (
2 2 2
= + + ÷ + + + ÷ +       v b b b v (5)




Numerical Analysis

Table 1. Numerical Analysis of Stability Equilibrium Point in FitzHugh-Nagumo Systems

No Variation I
eks
Equilibrium Point Eigen value Stability
1 0.00 -1.1994,-0.6243 -0.2513 ± 0.211900 i Spiral sink
2 0.32 -0.9769,-0.3461 -0.009176 ± 0.2774 i Spiral sink
3 0.33 -0.9685,-0.3357 -0.001045 ± 0.2757 i Limit cycle
4 0.50 -0.1311,-0.8048 +0.1441 ± 0.191500 i Spiral source
5 1.25 1.8810, 0.8048 +0.1441 ± 0.191500 i Spiral source
6 1.42 2.0857, 0.9685 -0.001045 ± 0.2757 i Spiral sink
7 1.43 0.9769, 2.0961 -0.009176 ± 0.2774 i Spiral sink
8 1.45 0.9933, 2.1166 -0.02532 ± 0.28020 i Spiral sink
9 1.50 2.1656, 1.0325 -0.06501 ± 0.282800i Spiral sink
10 2.00 1.3341, 2.5426 -0.6412, -0.2026 Stable Node

Obviously, this figures show transitions from stable to periodic or quasiperiodic, then
complex and finaly chaotic behaviour.


(a) (b) (c) (d)
Figure 2. Phases portrait for; (a) I = 0, (b) I = 0.33, (c) I = 1.45, (d) I = 2



(e) (f) (g) (h)
Figure 3. Time Series for ; (a) I = 0, (b) I = 0.33, (c) I = 1.45, (d) I = 2









Synchronization

Generalities

Synchronization is a phenomenon
characteristic of many processes in natural
systems and non linear science. It has
remained an objective of intensive
research and is today considered as one of
the basic nonlinear phenomena studied in
mathematics, physics, engineering or life
science.
This word has a greek root, syn =
common and chronos = time, which
means to share common time or to occur
at the same time, that is correlation or
agreement in time of different
processes.Thus, synchronization of two
dynamical systems generally means that
one system somehow traces the motion of
another.
Chaotic oscillators are found in many
dynamical systems of various origins. The
behavior of such systems is characterized
by instability and, as a result, limited
predictability in time. Roughly speaking, a
system is chaotic if it is deterministic, if it
has a long-term aperiodic behavior, and if
it exhibits sensitive dependence on initial
conditions on a closed invariant set.
Consequently, for a chaotic system,
trajectories starting arbitrarily close to
each other diverge exponentially with
time, and quickly become uncorrelated. It
follows that two identical chaotic systems
cannot synchronize. This means that they
cannot produce identical chaotic signals,
unless they are initialized at exactly the
same point, which is in general physically
impossible. Thus, at first sight,
synchronization of chaotic systems seems
to be rather surprising because one may
intuitively expect that the sensitive
dependence on initial conditions would
lead to an immediate breakdown of any
synchronization of coupled chaotic
systems, which led to the belief that chaos
is uncontrollable and thus unusable.
Despite this, in the last decades, the
search for synchronization has moved to
chaotic systems. A lot of research has
been carried out and, as a result, showed
that two chaotic systems could be
synchronized by coupling them :
synchronization of chaos is actual and
chaos could then be exploited. Ever since,
many researchers have discussed the
theory, the design or applications of
synchronized motion in coupled chaotic
systems. A broad variety of applications
have emerged, for example to increase the
power of lasers, to synchronize the output
of electronic circuits, to control
oscillations in chemical reactions or to
encode electronic messages for secure
communications (Aziz-Alaoui, MA.,
2006).

Modified FitzHugh-Nagumo Models

The modified FitzHugh-Nagumo
equations are a set of two coupled
differential equations which exhibit the
qualitative behavior observed in neurons,
viz quiescence, excitability and periodicity
(Mishra D et al, 2006). The system can be
represented as

) (
) (
3
1
3
bw a v
dt
dw
t I w v v
dt
dv
÷ + =
+ ÷ ÷ =



Where

) cos( ) / ( ) ( t A t I O O = (6)
A represents the magnitude of the stimulus
and O refers to the frequency of given
stimulus.






6

(a) (b)

(c) (d)

(e) (f)
Figure 4. FitzHugh-Nagumo Models at A = 0.7 ; (a) Time series at O= 0.4 (b) Phases portrait at O= 0.4 , (c) Time
series at O= 0.8145 (d) Phases portrait at O= 0.8145 , (e) Time series at O= 0.93, (f) Phases portrait at O= 0.93

The stimulus frequency is varied while
keeping the magnitude at a fixed value of
A = 0.71, since at this particular value of
A, modified FitzHugh-Nagumo neuron
model gives periodic spiking. Simulation
results at different stimulus frequencies
are shown in Figure 4. It is observed that
with the variation in stimulus frequency,
the neuron shows complex chaotic
behavior. Hence the stimulus frequency
can be considered as a significant
parameter that affects the behavior of
neuron.







7

Coupling FitzHugh-Nagumo neurons

Let us consider a network composed
by n FitzHugh-Nagumo neurons. These
neurons are coupled by there first variable
v. This network can be modeled by the
system (Lange, E.et al, 2005; Belykh, I &
Shilnikov A., 2008):

) (
) , ( ) (
3
1
3
i i
i
j i i i i
i
bw a v
dt
dw
v v h t I w v v
dt
dv
÷ + =
÷ + ÷ ÷ =

(7)

The coupling function h is given by :

¿
=
I ÷ =
n
j
j ij s s i j i
v c g V v v v h
1
) ( ) ( ) , ( (8)

in which the synaptic coupling Γ is
modeled by a sigmoid function with a
threshold :

)) ( exp( 1
1
) (
s j
j
v
v
O ÷ ÷ +
= I

(9)

Θs is the threshold reached by every
action potential for a neuron. Neurons are
supposed to be identical and the synapses
are fast and instantaneous. The parameter
gs corresponds to the synaptic coupling
strength. The synapse is exitatory, that is
why the reversal potential Vs must be
larger than xi(t) for all i and all t.

Coupling two FitzHugh-Nagumo
neurons

The first step here is to adapt the
previous method to two FitzHugh-
Nagumo neurons. In this case, we use a
bidirectional connection since the number
of inputs (that is one) has to be the same
for all neurons to have a synchronous
solution.


(
¸
(

¸

=
0 1
1 0
2
C



The following system represents our two
FitzHugh-Nagumo neurons bidirectionally
coupled :

) (
)) ( exp( 1
1
) (
3
1
) (
)) ( exp( 1
1
) (
3
1
2 2
2
1
2 2
3
2 2
2
1 1
1
2
1 1
3
1 1
1
bw a v
dt
dw
v
g V v I w v v
dt
dv
bw a v
dt
dw
v
g V v I w v v
dt
dv
s
s s
s
s s
÷ + =
O ÷ ÷ +
÷ ÷ + ÷ ÷ =
÷ + =
O ÷ ÷ +
÷ ÷ + ÷ ÷ =




(10)

Parameters are fixed as follows : a = 0.7, b
= 0.8,  = 0.08, V
s
= 1,
s
O = -0.25,  =1,
A = 0.71 and O= 0.95.








8

(a) (b)

(c) (d)
Figure 5. v
2
according to v
1
for the coupling strength; (a) g
s
= 0.1, (b) g
s
= 0.125, (c) g
s
= 0.1255 , dan
(d) g
s
= 0.15


We observe that the synchronization
numerically appears for a coupling
strength gs ≥ 0.15. This
phenomenon is also observed for the
variables w, for the same numerical values
of parameters.

Coupling three FitzHugh-Nagumo
neurons

In the case of three neurons, we also
use bidirectional connections since the
number of inputs (that are two) has to be
the same for all neurons to have a
synchronous solution.



(
(
(
¸
(

¸

=
1 0 0
0 1 0
0 0 1
3
C

The following system represents our three
FitzHugh-Nagumo neurons bidirectionally
coupled :














9
) (
)
)) ( exp( 1
1
)) ( exp( 1
1
( ) (
3
1
) (
)
)) ( exp( 1
1
)) ( exp( 1
1
( ) (
3
1
) (
)
)) ( exp( 1
1
)) ( exp( 1
1
( ) (
3
1
3 3
3
2 1
3 3
3
3 3
3
2 2
2
3 1
2 2
3
2 2
2
1 1
1
3 2
1 1
3
1 1
1
bw a v
dt
dw
v v
g V v I w v v
dt
dv
bw a v
dt
dw
v v
g V v I w v v
dt
dv
bw a v
dt
dw
v v
g V v I w v v
dt
dv
s s
s s
s s
s s
s s
s s
÷ + =
O ÷ ÷ +
+
O ÷ ÷ +
÷ ÷ + ÷ ÷ =
÷ + =
O ÷ ÷ +
+
O ÷ ÷ +
÷ ÷ + ÷ ÷ =
÷ + =
O ÷ ÷ +
+
O ÷ ÷ +
÷ ÷ + ÷ ÷ =

 

 

 
(11)


(a) (b)

(c) (d)
Figure 6. v
2
according to v
1
for the coupling strength; (a) g
s
= 0.01, (b) g
s
= 0.062, (c) g
s
= 0.064 ,
dan (d) g
s
= 0.069.

Synchronization numerically appears
for a coupling strength gs = 0.069. This
phenomenon is also observed for the
variables w, for the same numerical values
of parameters.


Coupling four FitzHugh-Nagumo
neurons
There are different possible ways for
coupling four neurons with the same
number of input from other neurons. Here,
we decided to connect each neuron to all
the others.






10

(a) (b)


(c) (d)

Figure 7. v
2
according to v
1
for the coupling strength; (a) g
s
= 0.015, (b) g
s
= 0.018, (c) g
s
= 0.019 ,
dan (d) g
s
= 0.0195.


Synchronization numerically appears
for a coupling strength gs = 0.0195. This
phenomenon is also observed for the
variables w, for the same numerical values
of parameters.
This is an emergent property which
comes from the collective dynamics of n
neurons. Moreover, as given in figure, as
the number of neurons n goes larger, the
synchronization threshold g gets smaller.

Conclusion

Since the birth of Hodgkin-Huxley
equation in neurophysiological modelling,
researchers have made considerable efort
to try to analyze this system in diferent
way. Extensive eforts have been made to
discover chaos in many physical and
biological systems including neural
systems. We have investigated
bifurcations observed in the FitzHugh-
Nagumo neuron model as a simplified
models. By calculating bifurcations with
changing I
ext
we have identified the
parameter regions in which the FitzHugh-
Nagumo neuron exhibits properties of
excitability and oscillation as an existence
of a couple of stable equilibrium points
and one stable limit cycle.
In this paper, the characteristics of two
dimensional modified FitzHugh-Nagumo
neuron model is studied. Dynamical
behavior of the modified FitzHugh-
Nagumo system under external electrical
stimulation is presented and it is verified
that the introduction of periodic
stimulation modifies the dynamics of





11
biological system by presenting the
dynamical behavior for the modified
FitzHugh-Nagumo system under external
electrical stimulation.
Synchronization phenomenon is
exhibited in a neuron network, the
transmission of information between these
neurons is optimal and the network can
develop consciousness. This is an
emergent property which comes from the
collective dynamics of n neurons.
Moreover as the number of neurons n goes
larger, the synchronization threshold g
gets smaller. Consciousness is more
important when the number of neurons is
larger. This phenomenon which can
describe many self-organized complex
systems, like earthquakes, linguistic,
urban systems.

References

[1]. Aziz-Alaoui, MA. (2006), Complex
emergent properties and chaos (De)
synchronization, Emergent
Properties in Natural and Artificial
Dynamical Systems. Heidelberg :
Springer p129-147.
[2]. Belykh, I., Shilnikov A. (2008),
When Weak Inhibition Synchronizes
Strongly Desynchronizing Networks
of Bursting Neurons. PRL 101,
078102.
[3]. Edelstein-Keshet, L., (1988),
Mathematical Models in Biology.
Random House, New York.
[4]. FitzHugh, R. (1961), Impulses and
Physiological state in theoretical
models of nerve membrane.
Biophysics Journal.,I,445-466.
[5]. Hodgkin, A. L., and Huxley, A. F.
(1952), A quantitative description of
membrane current and its application
to conduction and excitation in
nerve. J. Physiol.,117.500-544.
[6]. Lange, E., Belykh, I.,Hasler, M.
(2005), Synchronization of Bursting
Neurons: What matters in the
Network Topology. PRL 94,
188101.
[7]. Mishra D, Yadav A, Ray S, Kalra
PK. (2006), Controlling
Synchronization of Modified
FitzHugh-Nagumo Neurons Under
External Electrical Stimulation.
NeuroQuantology Issue 1 Page 50-
67.
[8]. Muratov, C. B. (2008), A
quantitative approximation scheme
for the traveling wave solutions in
the Hodgkin-Huxley model.
arXiv.org/abs/nlin/0209053v1.

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