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an active pulse transmission

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J. Nagumo

To electronically simulate an animal nerve axon, the authors made an active pulse transmission line

using tunnel diodes. The equation of propagation for this line is the same as that for a simplified model

of nerve membrane treated elsewhere. This line shapes the signal waveform peculiar to this line, smaller

signals are amplified, larger ones are attenuated, narrower ones are widened and those which are wider

are shrunk, all approaching the above mentioned specific waveform. In addition, this line has a certain

threshold value in respect to the signal height, and signals smaller than the threshold or noise is

eliminated in the course of transmission. Because of the above-mentioned shaping action and the

existence of a threshold, this line makes possible highly reliable pulse transmission, and will be useful for

various kinds of information processing systems.

Introduction

In the case of conventional pulse transmission lines, provided they are of great length, signals

suffer attenuation and distortion as they travel down the line, whereas an electric pulse signal which is

transmitted along an animal nerve axon suffers neither attenuation nor distortion, regardless of the

distance covered. A pulse transmission line with such qualities will be useful for pulse transmission in

communication systems, data processing systems, electronics computers, etc.

It is our purpose to realize such a pulse transmission line by simulating the animal nerve axon.

Excitation of Nerve Axon

A great deal of research on the electro-physiology of the nerve axon has already been made, the

most important being that of Hodgkin and Huxley. They have made a quantitative study of excitation of

a nerve axon and the propagation of the excitation, and have derived the Hodgkin-Huxley equations (H-

H equations) which describe the phenomena.

In the case of a space clamp, that is, in a case where the excitation of a nerve axon is spatially

uniform, the H-H equations describing the excitation of the nerve axon.

Where:

I = the membrane current density (a/cm

2

) [inward current positive].

V= the membrane voltage (mV) [difference from resting potential, depolarization negative],

m = the sodium activation (dimensionless) [varying between 0 and 1]

h = the sodium inactivation (dimensionless) [varying between 0 and 1]

n = the potassium activation (dimensionless) [varying between 0 and 1]

t = time (msec)

C

o

= the membrane capacitance (f/cm

2

)

g

Na

= 120, g

K

= 36, g

L

= (0.3 mmho/cm

2

)

V

Na

= -115, V

K

= 12, V

L

= -10.5988 (mV)

Several investigations of the H-H equations by the use of analog computers and digital

computers have appeared in recent literature. Nevertheless, it is still difficult to make an electronic

simulator of the H-H equations on account of their complexities.

The BVP Model

Recently, FitzHugh ingeniously simplified the H-H equations in case of a space clamp, making

use o an analog computer, and proposed the following BVP model (Bonhoeffer-van der Pol model)

Where a, b and c are constants satisfying the relations

The variables u,w and J in the BVP model correspond to the pair of variables (V, m), the pair of

variables (h, n) and I in the first equation, respectively.

Fig.1 (a) shows the trajectories on the (u, w) plane of the BVP model (2) in the case where J=0,

while Fig. 1(b) shows the trajectories on the (u*,w*)-plane of (1) in the case where I=0. Here,

Comparing Fig. 1(a) to Fig. 1 (b), it is observed that there is fairly good correspondence between

the H-H equations and the BVP model. We shall therefore try to simulate (2) instead of (1).

An Electronic Simulation of the BVP Model

Let us consider the two-terminal circuit of Fig. 2, where TD is a tunnel diode. From Kirchhoffs law

Fig. 1- The qualitative similarity of (a) and (b) suggests that the BVP model can be considered as

belonging to the same class of excitable systems as the Hodgkin-Huxley model.

where f(e) is a function, as shown in Fig. 3, which repr sen ts the voltage vs current characteristic of the

tunnel diode. .For simplicity, we assume

It is seen that (5) is reduced to (2).

Next, we shall examine the conditions in (3) for the constants a, b and c.

The first condition: 1>b>0 is equivalent to

The second condition: c

2

<b is equivalent to

The last condition: 1>a>1 2/3b is equivalent to

In Fig. 4.

The conditions in (7), (8) and (10) imply that in the case of the current clamp, j=0, the circuit

shown in Fig. 2 is nothing but the well-known monostable circuit of the tunnel diode. In particular the

condition in (8) means that the circuit oscillates spontaneously if the bias voltage is set at e

4

in Fig. 4. In

fact, if E

o

=e

4

, then a=0, and in the case of j=0, (2) becomes as follows:

Since c

2

>b, 1>b>0, c>0, this differential equation belongs to the Lienard type which represents self-

oscillations.

Therefore it is concluded that the BVP model can be simulated by the monostable circuit shown.

However, condition (10) which restricts the bias voltage E

0

is too severe; since even if the bias voltage is

set as

In Fig. 4, the circuit in Fig. 2 remains monostable. The condition in (12) is written as

And hence we shall employ the

conditions

instead of (3), for the constants in the BVP model.

Propagation of Excitation

The equations which describe the propagation of the excitation along the nerve axon are easily

obtained from the equations of the space clamp.

Denoting the distance along the nerve axon by S (cm), the radius of the nerve axon by R

o

(cm)

and the specific resistance of the axosplasm by r

o

(Kohmcm), we find that

Consequently, taking (1) into account, the H-H partial differentiation equations describing the

propagation of the excitation become as follows:

Similarly, let us consider the circuit shown in Fig. 5(a) or Fig. 5(b) which is constructed by

cascading the many two-terminal circuits of Fig. 2 through interstage coupling resistances.

Regarding the above circuit as a distributed line, we have

Corresponding to (14), provided the interstage coupling resistance per unit length of the line is r. From

(2) and (16), it follows that

Where h=p/r

The system of partial difference equations (17) is the distributed BVP model which may be

considered as a simplified system of the H-H partial differential equations (15).

By eliminating w from (17), a single partial differential equation for u is obtained as follows:

For simplicity, let us consider a case where R=0 (b=0). In such a case, (18) takes the form

Fig. 5 A distributed active line is obtained by cascading the two terminal circuits in Fig. 2 through

interstage coupling resistance. The circuit shown in (b) is equivalent to that in (a).

Fig. 6 Electronic model corresponding to (22).

It seems likely that the partial differential equation (22) is one of the simplest mathematical models of

the nerve axon. The structure of the electronic model which corresponds to (22) is shown in Fig. 6.

A Boundary-Value Problem

We now proceed to investigate the following boundary-value problem for the partial differential

equation (22)(Fig 7).

Some results of the numerical calculation using digital computer are given below. We chose =10 =0.1

and

A. Signal Amplification

The case of t

o

=3.0, z

o

=5.0 is shown in Fig. 8. In this case the signal is amplified in the course of

transmission.

B. Signal Attenuation

The case of t

o

=1.0, z

o

=20.0 is shown in Fig. 9. In this case the signal is attenuated in the course of

transmission.

C. Signal Elimination

The case t

0

=3.0, z

o

=3.0 is shown in Fig. 10. In this case the signal is eliminated in the course of

transmission.

From there results, it is expected that with respect to the signal height (there being an

asymptotic value and a threshold value).

- A signal with a height between the asymptotic value and the threshold value is amplified

during transmission,

- A signal higher than the asymptotic value is attenuated to the asymptotic value during

transmission,

- A signal lower than the threshold value is eliminated during transmission.

These actions may be called shaping with respect to signal height. In order to clarify the

situation, we shall try to seek for the asymptotic waveform.

Asymptotic Waveform

If the partial differential equation has a waveform, as its solution, which is transmitted along a

line without suffering distortion and with a constant velocity (say ), then the solution must be a

function of r=t-x/. In such a case the substitution

Reduces the partial differential equation to an ordinary differential equation:

To electronically simulate the distributed BVP model, which may be considered as a simplified

Hodgkin-Huxley model describing the propagation of the excitation of the nerve axon, we have made an

approximately distributer active pulse transmission line using tunnel diodes. The line behaves similarly

to the living axon, but the representation is gross.

The line is constructed by cascading many monostable circuits and consists essentially of the

following for parts:

- Transverse power voltage supply,

- Transverse inductance,

- Transverse tunnel diode,

- Longitudinal coupling resistance.

The active line shows the following characteristics in the transmission of signals:

- There is a certain threshold value in respect to the signal height, and signals below the threshold

are eliminated during transmission.

- The line shapes signal waveforms. Namely, there being a specific pulse-like waveform peculiar to

this line, signals above the threshold approach it during transmission.

- Since this line has a symmetrical structure, the signal transmission is bidirectional. Two signals

travelling in opposite directions from both ends vanish at the collision point.

On account of the existence of the threshold and the shaping action, this line makes possible

highly reliable pulse transmission, and will be useful for various kinds of information-processing

systems.

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