We have so far been discussing fibers without the thick myelin sheath. Unmyelinated fibers constitute about two-thirds of the fibers in the human body . . . Myelinated fibers are relatively large, with outer radii of 0.5 – 10 μm. They are wrapped with many layers of myelin between the nodes of Ranvier . . . In the myelinated region the conduction of the nerve impulse can be modeled by electrotonus because the conductance of the myelin sheath is independent of voltage. At each node a regenerative Hodgkin-Huxley-type (HH-type) conductance change restores the shape of the pulse. Such conduction is called saltatory conduction because saltare is the Latin verb “to jump.”
If saltatory conduction is not the best description of propagation along a myelinated axon, then what is responsible for the speedup compared to unmyelinated axons? Primarily, the action potential propagates faster because of a reduction of the membrane capacitance. Along the myelinated section of the membrane, the capacitance is low because of the many layers of myelin (N capacitors C in series result in a total capacitance of C/N). At a node of Ranvier, the capacitance per unit area of the membrane is normal, but the area of the nodal membrane is small. Adding these two contributions together leads to a very small average, or effective, capacitance, which allows the membrane potential to increase very quickly, resulting in fast propagation.
In summary, I don’t find the idea of an action potential jumping from node to node to be the most useful image of propagation along a myelinated axon. Instead, I prefer to think of propagation as being nearly continuous, with the reduced effective capacitance increasing the speed. This isn’t the typical explanation found in physiology books, but I believe it’s closer to the truth. Rather than using the term saltatory conduction, I suggest we use curretory conduction, for the Latin verb currere, “to run.”
Very Nice! Just one example of how you have helped me 'see.'
ReplyDeleteI'm a week behind. Thanks for the post on a topic that is particularly important to me. I'll enjoy considering two this week.
ReplyDeleteThanks for link to '49 Huxley of which I am beginning a careful reading.
ReplyDeleteHis summary point #6,
"Conduction was blocked reversibly by increasing the external resistance between two adjacent nodes. It is concluded that the action potential at each node excites the next node by current flowing forward in the axis cylinder and back in the fluid outside the myelin sheath."
supports my contention that low intensity, pulsed, focused ultrasound can be employed to reversibly, noninvasively block nerve conduction.