Action Potential Propagation along Myelinated Axons

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An action potential travels down the myelinated axon of a neuron. In between myelin sheath sections (indicated by light blue boxes) at the nodes of Ranvier (indicated by vertical red lines in the plot), the potential is modeled by Hodgkin–Huxley dynamics. The propagation of the signal along the myelinated sections is described by the linear cable equation, whose space and time constants are variable, as well as by the inter-node distance. The signal is initiated at the left-end node by an external current pulse or constant step of adjustable magnitude. The time course of a single point along the axon is displayed in the top right. You can see the speed of signal propagation in real time using the autoplay feature of the time slider.

Contributed by: Oliver K. Ernst (July 2015)
Open content licensed under CC BY-NC-SA


Snapshots


Details

Snapshot 1: the action potential propagating down the axon at a sufficiently short inter-node length

Snapshot 2: at low-time constants and high-space constants, the cable becomes rigid

Snapshot 3: the external current may be a pulse or a constant step, with the latter case allowing for successive signals to propagate down the axon

The speed of signal propagation is of critical importance for information processing in the nervous system. Since the axons of neurons can extend to over one meter in length, biological mechanisms have evolved that increase the propagation velocity of the action potential.

The axons of typical neurons are ensheathed in layers of a dielectric material called myelin. The width of the sheath is approximately 30–40% of the total diameter of the axon, and is interrupted periodically by nodes, termed the nodes of Ranvier. The layers of myelin both decrease the capacitance of the membrane and increase its resistance, leading to an increase in the speed of signal propagation and a reduction of radial current leakage [1].

The present Demonstration follows a classical model of the membrane potential in myelinated axons proposed by Fitzhugh in 1962 [2].

At the nodes of Ranvier, the membrane potential is modeled by Hodgkin–Huxley (HH) dynamics, with parameters given in [4]. In the present work, since the length of the node is short (c. 0.1% [1]) relative to the myelinated section, the nodes are modeled by a single HH compartment with no spatial dependence. Along the myelin sheath sections, the absence of voltage-gated ionic channels allows for the potential to be described by the linear cable equation [2]. This may be written as

,

where is the membrane potential (outside minus inside), mV is the resting potential, is the space constant, and is the time constant, which may be expressed as

,

,

where is the membrane resistance per unit length, is the intracellular resistance per unit length along the cable, and is the capacitance per unit length. These are further related physical properties of the axon and the myelin sheath by

,

,

,

where the parameters are the specific intracellular resistivity , specific membrane resistance , specific membrane capacitance , and fiber diameter [1].

An external current is applied at the cell body end of the axon [4] as either a pulse or a constant step of adjustable magnitude.

The boundary conditions between the node and inter-node dynamics are given explicitly in [2].

You can directly control the space and time constants, which in turn represent changes in the physical property of the myelin sheath. Additionally, you can vary the length of the myelin segments and the strength of the stimulating currents.

Using the autoplay feature, you can observe the speed of signal propagation in real time. Anatomically it has been observed that the inter-node distance and thus the velocity of the signal is proportional to the diameter of the axon [1]. Therefore, the speed may be increased by increasing the space constant or decreasing the time constant given above.

[1] C. Koch, Biophysics of Computation: Information Processing in Single Neurons, New York: Oxford University Press, 1999.

[2] R. Fitzhugh, "Computation of Impulse Initiation and Saltatory Conduction in a Myelinated Nerve Fiber," Biophysical Journal, 2(1), 1962 pp. 11–21.

[3] A. L. Hodgkin and A. F. Huxley, "A Quantitative Description of Membrane Current and Its Application to Conduction and Excitation in Nerve," The Journal of Physiology, 117(4), 1952 pp. 500–544.



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