CAUCHY – JURNAL MATEMATIKA MURNI DAN APLIKASI

Volume ##(##) (####), Pages 1-11

p-ISSN: 2086-0382; e-ISSN: 2477-3344

Analysis of the Hodgkin-Huxley Model on Transmission of Nerve Cell

Impulse Action Potentials

Mukhammad Fahmi1, Usman Pagalay1, Imam Sujarwo1

1Universitas Islam Negeri Maulana Malik Ibrahim Malang

Email: mukhammadfahmi25@gmail.com, usmanpagalay@yahoo.co.id

imamsoejarwo@gmail.com

ABSTRACT
The Hodgkin-Huxley model explains that on the Neuron membrane there are special channels
that only can be accessed by certain ions, namely Potassium (channel for K ions), Sodium
(channel for Na ions), and Leakage channel (channel for Leakage ions) which can lead to the
transmission of nerve cell impulse of action potentials. The Hodgkin-Huxley model consists of
four ordinary differential equations dependent of time, one being differential equations for the
membrane potentials of Neuron and the other are differential equations for the ion population
inside the gate which are the main features of the Neuron membrane of channel mechanism. The
results of the research indicate that the Hodgkin-Huxley model is expressed saddle with unstable
stability because 𝜆2 < 0 < 𝜆1 and the multiplication of 𝜆1 ∙ 𝜆2 is negative. Then the complex part
is expressed spiral with asymptotically stable because 𝜆3,4 = 𝑎 + 𝑏𝑖 ∈ 𝐶 with 𝑎 < 0. The result of
numerical simulation using ODE45 obtained that Hodgkin-Huxley model becomes unstable
when the external current at the interval of 0.1 𝐴 cm−2 < 𝐼ext < 2 𝐴 cm−2 . While at the interval
𝐼ext ≥ 2 𝐴 cm−2 the graph is stable and goes in the direction of the equilibrium point.

Keywords: Ionic Current, Neuron Membrane Potentials, the Hodgkin-Huxley Model.

INTRODUCTION
Allah Swt completes the incidence of Neuron membranes with the availability of
Sodium ions and Potassium ions in a balanced manner in the human body. In the nerve
cell system there is a membrane potential of Neuron 𝑉(𝑡) which acts as an impulse
receiver in the form of an electric current of the signal carrier. This signal will activate
the spread 1st level of Na ions, 2nd level of Na ions, K ions, and Leakage ions. The ions will
pass through special channels. The 1st level of Na ions passes through Sodium (the
channel for the Na ions) which then enters the 𝑚(𝑡) holding gate. The 2nd level of Na
ions will also pass through Sodium, which then enters the ℎ(𝑡) holding gate. The K ions
passes through Potassium (the channel for Potassium ions) which then enters the 𝑛(𝑡)
holding gate. The Leakage ion passes through the Leakage ion channel and goes directly
to the outside of the Neuron membrane, as there is no Leakage ion-holding gate.
 Analysis of the Hodgkin-Huxley Model on Transmission of Nerve Cell Impulse Action Potentials

In 1952, Alan Lloyd Hodgkin and Andrew Huxley introduced the Hodgkin-Huxley
model. The Hodgkin-Huxley model explains that on the Neuron membrane there are
special channels that can only be accessed by certain ions, namely Potassium (channel
for K ions), Sodium (channel for Na ions), and Leakage channel (channel for Leakage
ions) which can lead to the transmission of nerve cell impulse action potentials. Previous
research has been conducted by Netanel Ofer and Orit Shefi (2016). They studied the
four-variable Hodgkin-Huxley cables model by looking at the axon geometry side and
the Neuron membrane capacitance (Ofer & Shefi, 2016).
The problem raised in this research is how to know the Hodgkin-Huxley model
analysis to describe the model when there is a change in electric current impulse 𝐼ext
received by the membrane potential 𝑉(𝑡) from outside the Neuron membrane or in
other words, when the value of 𝐼ext value is fickle. By observe at this change of 𝐼ext value
is expected to control the magnitude of Neuron membrane potential and the ionic
population of Neuron membrane. The numerical simulation conducted in this research
is by using the method of Heun, Runge Kutta, and ODE45. So the dynamics obtained from
changes in the ionic population is controlled by the magnitude of the membrane
potential. Thus, the depth of this research is to see how far the Hodgkin-Huxley model
analysis of the external current changes. Therefore, it can be concluded how the
condition of an ionic population of Neuron membranes to remain balanced.

LITERATURE REVIEW
2.1 The Hodgkin-Huxley Model
The Hodgkin-Huxley model consists of four ordinary differential equations that
dependent of time, one being differential equations for the membrane potential of
Neuron and the other three are differential equations for the ion population inside the
gate which are the main features of the Neuron membrane channel mechanism.
The Hodgkin-Huxley model consists of independent variables, bound variables,
and several parameters. The independent variable is time t, the dependent variable is
the membrane potential of 𝑉(𝑡), the population 1st level of Na ions 𝑚(𝑡), the population
K ions 𝑛(𝑡), and the population 2nd level of Na ions ℎ(𝑡). The parameters are 𝐶M
membrane capacitance, the maximum conductance of Sodium 𝑔̅Na ions, the maximum
conductance of Potassium 𝑔̅K ions, the maximum conductance of Leakage 𝑔̅L ions,
Sodium ions resting potential 𝐸Na , Potassium ions resting potential 𝐸K , and Leakage ion
resting potential 𝐸L .

2.2 Differential Equations, Vector Fields, and Linear Algebra

Natural, social and artificial systems change hour by hour. Dynamical system is a
mathematics for the modeling and the analysis of such systems’ behavior. Dynamical
systems incorporate the state and its time change in a system. Consider a difference
equation or continuous-time dynamical system (Doi, Inoue, Pan, & Tsumoto, 2010).
𝑑 (2.1)
𝑥(𝑡) = 𝑓(𝑥(𝑡)), 𝑥(𝑡) ∈ ℝ𝑁 , 𝑡∈ℝ
𝑑𝑡
The solution 𝑥(𝑡) which satisfies this differential equation is called an orbit or trajectory
of the system (2.1). The state 𝑥(0) is referred as the initial state again. The special state
point 𝑥 ∗ such that 𝑓(𝑥 ∗ ) = 0 is called a fixed point, an equilibrium point or a steady state.
The right-hand side (r.h.s.) 𝑓(𝑥(𝑡)) ∈ ℝ𝑁 of the differential equation (2.1) is a vector and
is called a vector field. The vector field assigns the vector 𝑓(𝑥) to each point 𝑥 of the state

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 Analysis of the Hodgkin-Huxley Model on Transmission of Nerve Cell Impulse Action Potentials

space ℝ𝑁 . The simplest example of the continuous dynamical system (2.1) is also the
case that the map 𝑓 is a linear matrix 𝐴:
𝑑 (2.2)
𝑥(𝑡) = 𝐴𝑥(𝑡), 𝑥(𝑡) ∈ ℝ𝑁 , 𝑡∈ℝ
𝑑𝑡
where A is an invertible 𝑁 × 𝑁 matrix. Note that only the origin 𝑥 = (0, … ,0)𝑇 is the
fixed point or an equilibrium point since this system is linear and 𝐴 is invertible. The
general solutions can be obtained by
𝑥(𝑡) = exp(𝐴𝑡)𝑥(0) (2.3)
A vector 𝑥 ≠ 0 is called an eigenvector of a matrix 𝐴 if the following equation
holds
𝐴𝑥 = 𝜆𝑥 (2.4)
where the number 𝜆 is called an eigenvalue of 𝐴. Let the eigenvalue and the eigenvector
of a matrix 𝐴 be 𝜆 and 𝑣. Then, in general, the orbit with an initial value which is multiple
of 𝑣 becomes
𝑥(𝑛) = 𝐴𝑛 (𝛼𝑣) = 𝛼𝜆𝑛 𝑣 (2.5)
Thus, all orbits whose initial values (vectors) are scalar-multiples of eigenvector move
straightly whereas other orbits move curvedly.

2.3 Linearization and Stabilities

In the case of the continuous-time dynamical system (2.1), the equilibrium point
at the origin is stable if the real parts of all eigenvalues of 𝐴 are less than zero (negative).
Next, consider nonlinear dynamical systems. Let 𝑥 ∗ be an equilibrium point of the
continuous-time dynamical system (2.1), then 𝑓(𝑥 ∗ ) = 0 in (2.1). The Taylor expansion
of the function 𝑓(𝑥) near the equilibrium point 𝑥 ∗ are obtained as follows (Doi et al.,
2010):
2
𝑓(𝑥) = 𝑓(𝑥 ∗ ) + 𝒟𝑓(𝑥 ∗ )(𝑥 − 𝑥 ∗ ) + 𝒪 (||𝑥 − 𝑥 ∗ || ) (2.6)
2
where (||𝑥 − 𝑥 ∗ || ) denotes the higher-order terms (second-order terms and higher
terms) and 𝒟𝑓(𝑥 ∗ ) is the Jacobian matrix:
𝜕𝑓1 (𝑥) 𝜕𝑓1 (𝑥) (2.7)
⋯
𝜕𝑥1 𝜕𝑥2
𝒟𝑓(𝑥 ∗ ) = 𝜕𝑓2 (𝑥) 𝜕𝑓2 (𝑥)
⋯
𝜕𝑥1 𝜕𝑥2
( ⋮ ⋮ ⋯ )𝑥 = 𝑥 ∗
𝑥 = (𝑥1 , 𝑥2 , ⋯ )𝑇 , 𝑓(𝑥) = (𝑓1 (𝑥1 , 𝑥2 , ⋯ ), 𝑓2 (𝑥1 , 𝑥2 , ⋯ ), ⋯ )𝑇
Near the equilibrium point 𝑥 ∗ , we can neglect (under some conditions) the higher-order
2 2 2
terms 𝒪 (||𝑥 − 𝑥 ∗ || ) since 𝒪 (||𝑥 − 𝑥 ∗ || ) becomes small when ||𝑥 − 𝑥 ∗ || is small.
Then, we can obtain a linearized system or linearization of (2.1), respectively as follows:
𝑑 (2.18)
𝑧(𝑡) = 𝐴𝑧(𝑡), 𝐴 = 𝒟𝑓(𝑥 ∗ )
𝑑𝑡
where we have made use of the change of a variable 𝑧(𝑡) = 𝑥(𝑡) − 𝑥 ∗ .

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 Analysis of the Hodgkin-Huxley Model on Transmission of Nerve Cell Impulse Action Potentials

RESULTS AND DISCUSSION

3.1. Analysis of the Hodgkin-Huxley Model
3.1.1 Identification of Variables and Parameters
The model used in this research is a system of four variables Hodgkin-Huxley
model that created by Hodgkin and Huxley. The equation is as follows (A. L. Hodgkin and
A. F. Huxley, 1952):

𝑑𝑉(𝑡) −𝐺(𝑉(𝑡), 𝑚(𝑡), 𝑛(𝑡), ℎ(𝑡)) + 𝐼ext (3.1𝑎)

=
𝑑𝑡 𝐶M
𝑑𝑚(𝑡) (3.1𝑏)
= 𝛼𝑚 (𝑉(𝑡))(1 − 𝑚(𝑡)) − 𝛽𝑚 (𝑉(𝑡))𝑚(𝑡)
𝑑𝑡
𝑑𝑛(𝑡) (3.1𝑐)
= 𝛼𝑛 (𝑉(𝑡))(1 − 𝑛(𝑡)) − 𝛽𝑛 (𝑉(𝑡))𝑛(𝑡)
𝑑𝑡
𝑑ℎ(𝑡) (3.1𝑑)
= 𝛼ℎ (𝑉(𝑡))(1 − ℎ(𝑡)) − 𝛽ℎ (𝑉(𝑡))ℎ(𝑡)
𝑑𝑡
where
𝐺(𝑉(𝑡), 𝑚(𝑡), 𝑛(𝑡), ℎ(𝑡)) = 𝐼Na (𝑉(𝑡), 𝑚(𝑡), ℎ(𝑡)) + 𝐼K (𝑉(𝑡), 𝑛(𝑡)) + 𝐼L (𝑉(𝑡))
3 4
= 𝑔̅Na (𝑚(𝑡)) ℎ(𝑡)(𝑉(𝑡) − 𝐸Na ) + 𝑔̅K (𝑛(𝑡)) (𝑉(𝑡) − 𝐸K ) +
𝑔̅L (𝑉(𝑡) − 𝐸L )
with
0.1(𝑉(𝑡) + 35) 𝛽𝑚 (𝑉(𝑡)) = 4𝑒 (−0.0556(𝑉(𝑡)+60))
𝛼𝑚 (𝑉(𝑡)) = −(𝑉(𝑡)+35)
(1 − 𝑒 10 )
0.01(𝑉(𝑡) + 50) −(𝑉(𝑡)+60)
𝛼𝑛 (𝑉(𝑡)) = −(𝑉(𝑡)+50)
𝛽𝑛 (𝑉(𝑡)) = 0.125𝑒 80

(1 − 𝑒 10 )
𝛼ℎ (𝑉(𝑡)) = 0.07𝑒 (−0.05(𝑉(𝑡)+60)) 1
𝛽ℎ (𝑉(𝑡)) =
1+ 𝑒 −0.1(𝑉(𝑡)+30)
The description initial value of variables and parameters that used in the system
of Hodgkin-Huxley model (3.1) can be seen in the appendix.

3.1.2 Interpretation of Hodgkin-Huxley Model

Figure 3.1 The Interaction Scheme of Variable and Parameters of Hodgkin-Huxley Model

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 Analysis of the Hodgkin-Huxley Model on Transmission of Nerve Cell Impulse Action Potentials

An interpretation of the Hodgkin-Huxley model that consisting of the membrane

potential of Neuron 𝑉(𝑡), the number population 1st level of Na ions 𝑚(𝑡), the number
population K ions, and the number population 2nd level of Na ions ℎ(𝑡) as follows.

𝑑𝑉(𝑡) −𝐺(𝑉(𝑡), 𝑚(𝑡), 𝑛(𝑡), ℎ(𝑡)) + 𝐼ext

=
𝑑𝑡 𝐶M
The change of the Neuron membrane potential 𝑉(𝑡) with respect to time 𝑡 is
influenced by the membrane capacitance 𝐶M , the external current 𝐼ext and the ionic
current 𝐺(𝑉(𝑡), 𝑚(𝑡), 𝑛(𝑡), ℎ(𝑡)). This membrane potential 𝑉(𝑡) releases the ionic
currents 𝐺(𝑉(𝑡), 𝑚(𝑡), 𝑛(𝑡), ℎ(𝑡)) from the inside of Neuron membrane and receives the
external currents 𝐼ext from the outside of Neuron membrane. The ionic currents of
𝐺(𝑉(𝑡), 𝑚(𝑡), 𝑛(𝑡), ℎ(𝑡)) that consist of Sodium ion currents 𝐼Na , Potassium ion currents
𝐼K , and Leakage ion currents 𝐼L . Then this membrane potential 𝑉(𝑡) releases the 1st level
of Na ions to the 𝑚(𝑡) gate that expressed as 𝛼𝑚 (𝑉(𝑡))(1 − 𝑚(𝑡)). The membrane
potential 𝑉(𝑡) also releases the 2nd level of Na ions to the ℎ(𝑡) gate that expressed as
𝛼ℎ (𝑉(𝑡))(1 − ℎ(𝑡)). Next, potential membrane 𝑉(𝑡) releases K ions to the 𝑛(𝑡) gate that
expressed as 𝛼𝑛 (𝑉(𝑡))(1 − 𝑛(𝑡)).

𝑑𝑚(𝑡)
= 𝛼𝑚 (𝑉(𝑡))(1 − 𝑚(𝑡)) − 𝛽𝑚 (𝑉(𝑡))𝑚(𝑡)
𝑑𝑡
The change of the number population the 1st level of Na ions 𝑚(𝑡) with respect to
time 𝑡 is affected by activation 1st level of Na ions to enter the 𝑚(𝑡) gate reduced by the
release of 1st level of Na ions. When the membrane potential 𝑉(𝑡) receives an external
current impulse 𝐼ext from outside the Neuron membrane, then the 1st level of Na ions
that inside of the Neuron membrane will pass through Sodium (the channel for the Na
ions) and enter the empty 𝑚(𝑡) gate. Then the 𝑚(𝑡) gate performs the activation and
receives the 1st level of Na ions that expressed as 𝛼𝑚 (𝑉(𝑡))(1 − 𝑚(𝑡)) and processes
them for some time. 𝛼𝑚 (𝑉(𝑡)) is the velocity of a the 1st level of Na ions to enter the
𝑚(𝑡) gate. While (1 − 𝑚(𝑡)) is the condition of the 𝑚(𝑡) gate which does not yet contain
of the population 1st level of Na ions. After processing, the 𝑚(𝑡) gate opens and releases
the outcome of the process to the outside of the Neuron membrane that expressed as
𝛽𝑚 (𝑉(𝑡))𝑚(𝑡). 𝛽𝑚 (𝑉(𝑡)) is the velocity 1st level of Na ions to exit the 𝑚(𝑡) gate. While
𝑚(𝑡) describes a gate containing population 1st level of Na ions.

Figure 3.2 Ionic Channel Gate Mechanism of Neuron Membrane

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 Analysis of the Hodgkin-Huxley Model on Transmission of Nerve Cell Impulse Action Potentials

𝑑𝑛(𝑡)
= 𝛼𝑛 (𝑉(𝑡))(1 − 𝑛(𝑡)) − 𝛽𝑛 (𝑉(𝑡))𝑛(𝑡)
𝑑𝑡
The change of the number population the K ions 𝑛(𝑡) with respect to time 𝑡 is
affected by activation K ions to enter the 𝑛(𝑡) gate reduced by the release of K ions.
When the membrane potential 𝑉(𝑡) receives an external current impulse 𝐼ext from
outside the Neuron membrane, then the K ions that inside of the Neuron membrane will
pass through Potassium (the channel for the K ions) and enter the empty 𝑛(𝑡) gate. Then
the 𝑛(𝑡) gate performs the activation and receives the K ions that expressed as
𝛼𝑛 (𝑉(𝑡))(1 − 𝑛(𝑡)) and processes them for some time. 𝛼𝑛 (𝑉(𝑡)) is the velocity of a the
K ions to enter the 𝑛(𝑡) gate. While (1 − 𝑛(𝑡)) is the condition of the 𝑛(𝑡) gate which
does not yet contain of the population K ions. After processing, the 𝑛(𝑡) gate opens and
releases the outcome of the process to the outside of the Neuron membrane that
expressed as 𝛽𝑛 (𝑉(𝑡))𝑛(𝑡). 𝛽𝑛 (𝑉(𝑡)) is the velocity K ions to exit the 𝑛(𝑡) gate. While
𝑛(𝑡) describes a gate containing population K ions.

𝑑ℎ(𝑡)
= 𝛼ℎ (𝑉(𝑡))(1 − ℎ(𝑡)) − 𝛽ℎ (𝑉(𝑡))ℎ(𝑡)
𝑑𝑡
The change of the number population the 2nd level of Na ions ℎ(𝑡) with respect to
time 𝑡 is affected by activation 2nd level of Na ions to enter the ℎ(𝑡) gate reduced by the
release of 2nd level of Na ions. When the membrane potential 𝑉(𝑡) receives an external
current impulse 𝐼ext from outside the Neuron membrane, then the 2nd level of Na ions
that inside of the Neuron membrane will pass through Sodium (the channel for the Na
ions) and enter the empty ℎ(𝑡) gate. Then the ℎ(𝑡) gate performs the activation and
receives the 2nd level of Na ions that expressed as 𝛼ℎ (𝑉(𝑡))(1 − ℎ(𝑡)) and processes
them for some time. 𝛼ℎ (𝑉(𝑡)) is the velocity of a the 2nd level of Na ions to enter the ℎ(𝑡)
gate. While (1 − ℎ(𝑡)) is the condition of the ℎ(𝑡) gate which does not yet contain of the
population 2nd level of Na ions. After processing, the ℎ(𝑡) gate opens and releases the
outcome of the process to the outside of the Neuron membrane that expressed as
𝛽ℎ (𝑉(𝑡))ℎ(𝑡). 𝛽ℎ (𝑉(𝑡)) is the velocity 2nd level of Na ions to exit the ℎ(𝑡) gate. While
ℎ(𝑡) describes a gate containing population 2nd level of Na ions.

3.1.3 Stability of the Hodgkin-Huxley Model

3.1.3.1 The Equilibrium Point of the System
𝑑𝑉 𝑑𝑚 𝑑𝑛
The equilibrium point of the system (3.1) is obtained if = 0, = 0, 𝑑𝑡 = 0
𝑑𝑡 𝑑𝑡
𝑑ℎ
and 𝑑𝑡 = 0. From the system of equations (3.1) is sought the equilibrium point value
with the help of the Matlab program obtained 𝑉 ∗ = −54.61923 mV, 𝑚∗ = 0.09765 cell
ml−1 , 𝑛∗ = 0.40234 cell ml−1 , dan ℎ∗ = 0.40503 cell ml−1 .

3.1.3.2 Linearization
The approach of the linear system around the equilibrium point 𝑉 ∗ , 𝑚∗ , 𝑛∗ , and ℎ∗
using the Taylor series and cut to first order. After a long process obtained the following
linear equations:
𝑑𝑉
= −1.28862𝑉 + 152.64875𝑚 − 164.32306𝑛 + 12.26747ℎ + 0.1 (3.2𝑎)
𝑑𝑡
𝑑𝑚
= 0.03503𝑉 − 3.28667𝑚 (3.2𝑏)
𝑑𝑡

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 Analysis of the Hodgkin-Huxley Model on Transmission of Nerve Cell Impulse Action Potentials

𝑑𝑛
= 0.00312𝑉 − 0.19554𝑛 (3.2𝑐)
𝑑𝑡
𝑑ℎ
= −0.00452𝑉 − 0.13206ℎ (3.2𝑑)
𝑑𝑡
3.1.3.3 Eigenvalues
The Eigen value is obtained by completing det(λI − J) = 0. With the help of
Matlab program obtained Eigenvalues 𝜆1 = 0.30055, 𝜆2 = −0.14823, 𝜆3 = − 2.12839𝐼
− 0.78650, 𝜆4 = 2.12839𝐼 − 0.78650.

3.1.3.4 Stability of Equilibrium Point

At the state of equilibrium point 𝑉 ∗ = −54.61923 mV, 𝑚∗ = 0.09765 cell ml−1,
𝑛 = 0.40234 cell ml−1 , and ℎ∗ = 0.40503 cell ml−1 the Hodgkin-Huxley model is
∗

expressed saddle with unstable stability because 𝜆2 < 0 < 𝜆1 and the multiplication of
𝜆1 ∙ 𝜆2 is negative. Then the complex part is expressed spiral with asymptotically stable
because 𝜆3,4 = 𝑎 + 𝑏𝑖 ∈ 𝐶 with 𝑎 < 0.

3.1.4 Phase Plane Analysis

The Hodgkin-Huxley model has four variables that are difficult to observe as a
whole in the phase field directly. Then the separation of four variables into sub-variables
to solve them. In the sub-system 𝑉(𝑡) − 𝑚(𝑡), 𝑉(𝑡) and 𝑚(𝑡) are dynamic variables
while 𝑛(𝑡) and ℎ(𝑡) are defined as values corresponding to parameters. In this condition,
𝑛(𝑡) and ℎ(𝑡) is defined as a parameter value, i.e. 𝑛 = 0.317677 and ℎ = 0.596120
obtained the result of phase field of sub-system 𝑉(𝑡) − 𝑚(𝑡) using pplane8.m program
in Matlab (Wallisch & et. al, 2009). There are three equilibrium points, namely 𝐸1∗ , 𝐸2∗ ,
and 𝐸3∗ . Then, obtained picture of phase field of sub-system 𝑉(𝑡) − 𝑚(𝑡) as follows.

V ' = ((1/Cm) (I - (120 m3 h (V - 115) + 36 n4 (V + 12) + 0.3 (V - 10.599)))) I = 0.1 h = 0.596120 V ' = ((1/Cm) (I - (120 m3 h (V - 115) + 36 n4 (V + 12) + 0.3 (V - 10.599)))) I = 0.1 h = 0.596120
m ' = (0.1 (25 - V))/(exp((25 - V) (1/10)) - 1) (1 - m) - 4 exp( - (1/18) V) m Cm = 1 n = 0.317677 m ' = (0.1 (25 - V))/(exp((25 - V) (1/10)) - 1) (1 - m) - 4 exp( - (1/18) V) m Cm = 1 n = 0.317677

1 0.1

0.9
𝐸3∗ 0.09

0.8 𝑑𝑚 0.08
=0
0.7
𝑑𝑡
0.07

0.6 0.06
𝐸2∗
𝐸1∗
m

0.5
m

0.05

0.4 0.04

0.3 0.03
𝑑𝑉
0.2 =0
𝑑𝑡 0.02
𝐸2∗
0.1 0.01

0 𝐸1∗ 0

0 20 40 60 80 100 120 -2 0 2 4 6 8 10
V V

Figure 3.3 (a) Phase Plane Sub-System 𝑉(𝑡) − 𝑚(𝑡) Hodgkin-Huxley Model with 𝑛 = 0.317677 and ℎ = 0.596120
(b) Enlargement of Phase Plane area of Sub-System 𝑉(𝑡) − 𝑚(𝑡)

We can see that 𝑉(𝑡) nullcline (curve purple) and 𝑚(𝑡) nullcline (orange curve)
intersect at three points 𝐸1∗ = (0.4747, 0.055971), 𝐸2∗ = (2.2384, 0.068672), and 𝐸3∗ =
(113.9201, 0.9992) which is the equilibrium point of the sub-system 𝑉(𝑡) − 𝑚(𝑡). Thus,
the transmission process of the action potential of nerve cell impulse proceeds as
follows: silence → depolarization → decrease ℎ(𝑡) → increase 𝑛(𝑡) → repolarization.
Equilibrium point 𝐸1∗ = (0.4747, 0.055971) with eigenvalues 𝜆1 = −0.15428 and 𝜆2 =
−4.6518 are declared nodes with asymptotically stable stability because 𝜆2 , 𝜆1 < 0. The

Mukhammad Fahmi 7
 Analysis of the Hodgkin-Huxley Model on Transmission of Nerve Cell Impulse Action Potentials

equilibrium point 𝐸2∗ = (2.2384, 0.068672) with the eigenvalues 𝜆1 = 0.16989 dan 𝜆2 =
−4.6524 is expressed saddle with unstable stability because 𝜆2 < 0 < 𝜆1 and the
multiplication of 𝜆1 ∙ 𝜆2 is negative. While equilibrium point 𝐸3∗ = (113.9201, 0.9992)
with eigenvalues 𝜆1 = −72.0309 dan 𝜆2 = −8.8986 are stated nodes with
asymptotically stable stability properties because 𝜆2 , 𝜆1 < 0. If a large external current
𝐼ext is applied to the Neuron in a stationary state 𝐸1∗ , the equilibrium point moves to the
right beyond the stable manifold state of 𝐸2∗ , and then to 𝐸3∗ corresponding to the
depolarization state of the Neuron.

3.2 Numerical Simulation of Hodgkin-Huxley Model with Heun, Runge Kutta, and
ODE45 Methods

Figure 3.4 Comparison of Solutions 𝑉, 𝑚, 𝑛, ℎ with respect to 𝑡 with Heun, Runge Kutta, and ODE45 Methods

The Hodgkin-Huxley model has no analytical solution, so it can be used numerical

simulation with Heun, Runge Kutta, and ODE45 methods. The numerical method as an
alternative to the numerical solution of the analytic method because the method is a
meticulous method. It is said to be thorough as it shows that the resulting solution is in
line with the process of transmission of nerve cell impulse of action potentials. However,
the magnitude of the accuracy can not be measured, this is because the Hodgkin-Huxley
model is a nonlinear differential equation that can not be solved analytically or has no
exact solution. Because it has no exact solution, then the error can not be generated or
no true error.

3.2.1 When given 𝑰𝐞𝐱𝐭 = 𝟎. 𝟏 𝑨 𝐜𝐦−𝟐

Potensial Membran Neuron Model Hodgkin-Huxley Populasi Ionik Membran Neuron
60 1
V ODE45 m ODE45
0.9 n ODE45
40
h ODE45
0.8
Potensial Membran Neuron (mV)

Populasi Ionik Membran Neuron

20 0.7

0.6
0
0.5
-20
0.4

-40 0.3

0.2
-60
0.1

-80 0
0 5 10 15 20 25 30 35 40 45 50 0 5 10 15 20 25 30 35 40 45 50
Waktu (ms) Waktu (ms)

Figure 3.5 Graph of 𝑉, 𝑚, 𝑛, ℎ with respect to 𝑡 when given 𝐼ext = 0.1 𝐴 cm−2

Mukhammad Fahmi 8
 Analysis of the Hodgkin-Huxley Model on Transmission of Nerve Cell Impulse Action Potentials

Figure 3.5 is a graph of changes in the membrane potential of Neurons and

changes in Na, K, and Leakage ion populations that activate for 50 ms. When the neuron
membrane is given 𝐼ext = 0.1 𝐴 cm−2 then obtained the equilibrium point 𝑉 ∗ = −54.619
23 mV, 𝑚∗ = 0.09765 cell ml−1 , 𝑛∗ = 0.40234 cell ml−1 , dan ℎ∗ = 0.405 03 cell ml−1 .

3.2.2 When given 𝑰𝐞𝐱𝐭 = 𝟏 𝑨 𝐜𝐦−𝟐

Potensial Membran Neuron Ketika Iext=1 A/cm2 Populasi Ionik Membran Neuron Ketika Iext=1 A/cm2
60 1
V ODE45 m ODE45
0.9 n ODE45
40
h ODE45
0.8
Potensial Membran Neuron (mV)

Populasi Ionik Membran Neuron

20 0.7

0.6
0
0.5
-20
0.4

-40 0.3

0.2
-60
0.1

-80 0
0 5 10 15 20 25 30 35 40 45 50 0 5 10 15 20 25 30 35 40 45 50
Waktu (ms) Waktu (ms)

Figure 3.6 Graph of 𝑉, 𝑚, 𝑛, ℎ with Respect to 𝑡 when Given 𝐼ext = 1 𝐴 cm−2

When the neuron membrane is given 𝐼ext = 1 𝐴 cm−2 then obtained the
equilibrium point 𝑉 ∗ = −41.57391 mV, 𝑚∗ = 0.32994 cell ml−1 , 𝑛∗ = 0.59847 cell ml−1 ,
dan ℎ∗ = 0.10434 cell ml−1 .

3.2.3 When given 𝑰𝐞𝐱𝐭 = 𝟐 𝑨 𝐜𝐦−𝟐

Potensial Membran Neuron Ketika Iext=2 A/cm2 Populasi Ionik Membran Neuron Ketika Iext=2 A/cm2
60 1
V ODE45 m ODE45
0.9 n ODE45
40 h ODE45
0.8
Potensial Membran Neuron (mV)

Populasi Ionik Membran Neuron

0.7
20
0.6

0 0.5

0.4
-20
0.3

0.2
-40
0.1

-60 0
0 5 10 15 20 25 30 35 40 45 50 0 5 10 15 20 25 30 35 40 45 50
Waktu (ms) Waktu (ms)

Figure 3.7 Graph of 𝑉, 𝑚, 𝑛, ℎ with Respect to 𝑡 when Given 𝐼ext = 2 𝐴 cm−2

When the neuron membrane is given 𝐼ext = 2 𝐴 cm−2 then obtained the
equilibrium point 𝑉 ∗ = −35.84293 mV, 𝑚∗ = 0.47861 cell ml−1 , 𝑛∗ = 0.66919 cell ml−1 ,
dan ℎ∗ = 0.05521 cell ml−1 . The result of numerical simulation using ODE45 obtained
that Hodgkin-Huxley model becomes unstable when the external current at the interval
of 0.1 𝐴 cm−2 < 𝐼ext < 2 𝐴 cm−2. While at the interval 𝐼ext ≥ 2 𝐴 cm−2 the graph is stable
and goes in the direction of the equilibrium point.

Mukhammad Fahmi 9
 Analysis of the Hodgkin-Huxley Model on Transmission of Nerve Cell Impulse Action Potentials

CLOSING
4.1 Conclusion
Based on the result and discussion analysis of Hodgkin-Huxley model on the
transmission of nerve cell impulse action potential, it can be concluded that:
1. The Hodgkin-Huxley model is expressed saddle with unstable stability because 𝜆2 <
0 < 𝜆1 and the multiplication of 𝜆1 ∙ 𝜆2 is negative. Then the complex part is
expressed spiral with asymptotically stable because 𝜆3,4 = 𝑎 + 𝑏𝑖 ∈ 𝐶 with 𝑎 < 0. In
the phase-phase analysis, it is understood that the process of transmitting the nerve
cell impulse action potential in the 𝑉(𝑡) − 𝑚(𝑡) sub-system of Hodgkin-Huxley model
runs as follows: the resting state → depolarization → decrease of ℎ(𝑡) → increase of
𝑛(𝑡) → repolarization.
2. The numerical method as an alternative to the numerical solution of the analytic
method because the method is a meticulous method. It is said to be thorough as it
shows that the resulting solution is in line with the process of transmission of nerve
cell impulse of action potentials. However, the magnitude of the accuracy can not be
measured, this is because the Hodgkin-Huxley model is a nonlinear differential
equation that can not be solved analytically or has no exact solution. Because it has no
exact solution, then the error can not be generated or no true error. The result of
numerical simulation using ODE45 obtained that Hodgkin-Huxley model becomes
unstable when the external current at the interval of 0.1 𝐴 cm−2 < 𝐼ext < 2 𝐴 cm−2 .
While at the interval 𝐼ext ≥ 2 𝐴 cm−2 the graph is stable and goes in the direction of
the equilibrium point.

4.2 Suggestion
For further research, it is advisable to the reader to examine an analytical
solution from the Hodgkin-Huxley model and then compare the results with a numerical
solution to see how much approximation error of the numerical method.

REFERENCES

A. L. Hodgkin and A. F. Huxley. (1952). A Quantitative Description of Membrane Current

and its Application to Conduction and Excitation in Nerve. Journal Physiol, 117(1),
500–544.
Doi, S., Inoue, J., Pan, Z., & Tsumoto, K. (2010). Computational Electrophysiology
Dynamical Systems and Bifurcations. New York: Springer Tokyo Dordrecht
Heidelberg London.
E. M. Izhikevich. (2004). Dynamical Systems in Neuroscience The Geometry of Excitability
and Bursting. London: The MIT Press.
Ofer, N., & Shefi, O. (2016). Axonal geometry as a tool for modulating firing patterns.
Applied Mathematical Modelling, 40(1), 3175–3184.
Siciliano, R. (2012). The Hodgkin-Huxley model - Its extensions, analysis and numerics.,
41.
Wallisch, P., & et. al. (2009). Matlab for Neuroscientist. Elsevier (Vol. 53).

(E. M. Izhikevich, 2004)(Doi et al., 2010)(A. L. Hodgkin and A. F. Huxley, 1952)(Siciliano,

2012)(Ofer & Shefi, 2016)(Wallisch & et. al, 2009)

Mukhammad Fahmi 10
APPENDIX

Table 1. Initial Value of Variables (E. M. Izhikevich, 2004).

Initial
Variable Variable Description Unit
Value
𝑉(𝑡) The magnitude of the Neuron membrane potential -60 mV
𝑚(𝑡) The number of population 1st level of Na ions 0.06 cell ml−1
𝑛(𝑡) The number of population K ions 0.31 cell ml−1
ℎ(𝑡) The number of population 2nd level of Na ions 0.6 cell ml−1

Table 2. Initial Value of Parameter (Siciliano, 2012).

Para- Initial
Parameter Description Unit
meter Value
𝐼ext The amount of external current of neuron membrane 0.1 𝐴 cm−2
𝐶M Membrane capacitance 0.01 𝜇F cm−2
𝑔̅Na Maximum conductance of the Na ions 1.2 mS cm−2
𝑔̅K Maximum conductance of the K ions 0.36 mS cm−2
𝑔̅L Maximum conductance of the Leakage ions 0.003 mS cm−2
𝐸Na Resting potentials of Na ions 55.17 mV
𝐸K Resting potentials of K ions -72.14 mV
𝐸L Resting potentials of Leakage ions -49.42 mV

Table 3. Functions that Contain 𝑉(𝑡) (Siciliano, 2012).

Function Function Description Unit
𝛼𝑚 (𝑉(𝑡)) The velocity 1st level of Na ions to enter the 𝑚(𝑡) gate cell ms −1
𝛼𝑛 (𝑉(𝑡)) The velocity of K ions to enter the 𝑛(𝑡) gate cell ms −1
𝛼ℎ (𝑉(𝑡)) The velocity 2nd level of Na ions to enter the ℎ(𝑡) gate cell ms −1
𝛽𝑚 (𝑉(𝑡)) The velocity 1st level of Na ions to exit the 𝑚(𝑡) gate cell ms −1
𝛽𝑛 (𝑉(𝑡)) The velocity of K ions to exit the 𝑛(𝑡) gate cell ms −1
𝛽ℎ (𝑉(𝑡)) The velocity 2nd level of Na ions to exit the ℎ(𝑡) gate cell ms −1