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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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