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On the Propagation of Electric Pulse along an Axon

On the Propagation of Electric Pulse along an Axon

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Abstract

Considering longitudinal flow in the axon plasma and introducing the substantial derivative, a nonlinear differential equation describing signal propagation along with the axon fiber instead of the cable equation is established. In contrast to the cable equation, which gives solutions that distort the profile of the pulse, a nonlinear equation describes a traveling wave of an unchangeable profile. The speed of the electric pulse wave, w = 320√a m/s, changes proportionally to the square root of the axon fiber radius, a.

I. INTRODUCTION

Analogous to the wires that connect different points in an electric circuit, axons are responsible for the transmission of information over the nervous system. The axon, as a part of a cell, separates the internal medium from the external one with the plasma membrane and any perturbations of a membrane potential (signal) conducted along the axon. The membrane potential is the result of ionic gradient currents perpendicular to the membrane.

The following events characterize the transmission of a nerve impulse (Figure 1):

  • The resting potential describes the polarized state of a neuron ( −70mV ).

  • A graded potential is a change in the resting potential in the response to a stimulus, which causes Na+ or K+ gated channels to open. If Na+ channels open, the membrane depolarizes. If the K+ channels open, then positive potassium ions exit across the membrane and hyperpolarizes it (Figure 1). A graded potential occurs in cell bodies and dendrites and is a local event that does not travel from its origin.

Figure 1: Events that characterize the transmission of a nerve impulse
Figure 1: Events that characterize the transmission of a nerve impulse
  • Unlike a static graded potential, an action potential is dynamic and capable of traveling along with a nerve fiber. If a graded potential is sufficiently large, Na+ channels open, and Na+ on the outside of the membrane becomes depolarized (-70 to +30 mV). If the stimulus exceeds a certain threshold level ( ∼55 mV) - additional Na+ gates open, increasing the flow of Na+, causing an action potential, which stimulates neighboring Na+ gates to open. In this manner, the action potential travels down the length of the axon as opened Na+ gates stimulate neighboring Na+ gates to open...

  • The inflow of Na+, K+ channels open, allowing K+ movement out of the cell causes repolarization by restoring the original membrane polarization by Na+/K+ pumps in the cell membrane.

II. CABLE EQUATION

Nerve impulses flow in only one direction, accompanied by currents across and along the axon trunk. Current across the membrane is

Im=CmdVmdt,

where Vm=Vin−Vout is the membrane potential, Cm=kε02πaL/b is the capacity of an unmyelinated cylindrical axon with a length of L (=1m), a membrane thickness of b = 10 nm, and a radius of a = 2.5 μ m. The length of the myelinated section is 1.4 mm and the thickness is b = 2 μ m.

In turn, these ionic currents give rise to longitudinal currents that allow the regeneration of the membrane potential changes in the axon [1]:

Ii=−dVridx,

where x is the axis along the axon, ri=ρiπa2 is the resistance per unit length of the axon plasma.

The propagation of the action potential is described by the well-known cable differential equation [1-3]:

(2)τ∂V∂t=λ2∂2V∂x2+ΔV,

where λ=abρm/2ρi is the space constant and τ=kε0ρm is the time constant,

ΔV=Vpeak−Vrest=−70mV.

The evolution of any initial impulse V(x′,0) is given by the integral

(3)V(v,t)=ΔV{tτ+τ4πt∫0∞V(x′,0)ΔVλexp[−τ(x−x′)24tλ2]dx′}.

The nerve impulse describing by this equation changes its form soon during propagation along with the nerve fiber. For example, for the evolution of δ -shape initial impulse, V(x′,0)=ΔVλδ(x′) integration leads to

(4)V(x,t)=ΔV[tτ+τ4πtexp(−τx24tλ2)],

that describes an ink drop diffusion in water, hence the nerve impulse spreads in space-time, distorting initial form, therefore the transmission information. Therefore, the transition of information through the nerve fiber should be occurring by a pulse-wave mechanism that does not change its initial shape.

III. A New Equation for the Signal Transmission

Because of longitudinal current, a flow of substance takes place along the axon plasma with an effective speed

u=−(ΣμiUiΣμi)∂V∂x,

where μi is the molar mass of ith type ion and Ui is corresponding mobility. The mass flow in the axon plasma requires the use of the substantial derivative,

ddt=∂∂t+u∂∂x,

instead of partial ∂∂t, in the left-hand side of the cable equation (2).

(5)τ∂V∂t−γλ2(∂V∂x)2=λ2∂2V∂x2+ΔV

where the constant parameter γ=τΣμiUi/λ2Σμi characterizes the longitudinal flow of substance.

Introduction of dimensionless running wave variable ξ=(x−wt)/λ, where w is the speed of transmission of an electric signal (potential change) over the nervous system, gives ordinary nonlinear differential equation for perturbed potential

(6)V′′+γV2+(wτλ)V′+ΔV=0,

where prime means derivative with respect ξ.

The general solution of nonlinear equation (4), considering w=λ/τ is:

V(ξ)=C2−ξ2γ+1γln⁡Cosh⁡121−4γΔV(ξ+C1)

For the intensity of electric field corresponding to generated potential (5), we obtain

(8)E(ξ)=−dVdx=12γ[1−1−4γΔVTanh121−4γΔV(ξ+C1)]

Taking into account the following boundary conditions:

V(0)=VpeakandV′(0)=0,

For constants C1 and C2 we obtain:

C1=21−4γΔVTanh−111−4γΔV,C2=V peak ΔV−1γΔVln⁡1−14γΔV.

With the use of numerical values of parameters [4,5], we can establish the profile of the electric intensity in the running wave (Fig. 2).

Figure 2: Profile of electric pulse propagation along an axon fiber
Figure 2: Profile of electric pulse propagation along an axon fiber

IV. CONCLUSION

Considering the longitudinal flow in the electrolyte of an axon plasma and introducing the substantial derivative, we have established nonlinear differential equation (3) describing signal propagation along with the axon fiber as a running wave without changing its profile in contrast to cable equation (2). The solution to this equation is obtained in the form of a running wave. The speed of this wave is

(10)w=λτ=abρiρmk2ε02=320ams

which changes proportionally to the square root of the axon fiber radius, a.

References

4 Cites in Article
  1. 1. Helmut Barz,Almut Schreiber,Ulrich Barz (2020). Nerve impulse propagation: Mechanical wave model and HH model. Medical Hypotheses, 137, 109540.
  2. 2. Martin Abrahamyan (2015). Application of Physics in Medicine. Application of Physics in Medicine, 263.
  3. 3. Debanjan Das,Farhan Kamil,Karabi Biswas,Soumen Das (2013). Evaluation of single cell electrical parameters from bioimpedance of a cell suspension. RSC Adv., 4(35), 18178-18185.
  4. 4. Yihao Zhang,Anastasios Tzingounis,George Lykotrafitis (2019). Modeling of the axon plasma membrane structure and its effects on protein diffusion. PLOS Computational Biology, 15(5), e1007003.

Funding

No external funding was declared for this work.

Conflict of Interest

The authors declare no conflict of interest.

Ethical Approval

No ethics committee approval was required for this article type.

Data Availability

Not applicable for this article.

How to Cite This Article

Martin G. Abrahamyan, Yanina Ya. Marinosyan. 2021. "On the Propagation of Electric Pulse along an Axon". Global Journal of Computer Science and Technology - G: Interdisciplinary GJCST-G Volume 21 (GJCST Volume 21 Issue G1).

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Journal Specifications

Crossref Journal DOI 10.17406/gjcst

Print ISSN 0975-4350

e-ISSN 0975-4172

Keywords
Classification
GJCST-G Classification G.1.0
Version of record
v1.2

Issue date
May 19, 2021

Language
English
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On the Propagation of Electric Pulse along an Axon

Martin Abrahamyan
Martin Abrahamyan
Yanina Marinosyan
Yanina Marinosyan